Results of the Archbold Frogs and Lizards from The

RESULTS OF THE ARCHBOLD EXPEDITIONS. NO. 103. FROGS AND LIZARDS FROM THE HUON PENINSUL.A, PAPUA NEW GUINEA

RICHARD G. ZWEIFEL

BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY VOLUME 165: ARTICLE 5 NEW YORK: 1980

RESULTS OF THE ARCHBOLD EXPEDITIONS. NO. 103. FROGS AND LIZARDS FROM THE HUON PENINSULA, PAPUA NEW GUINEA

RICHARD G. ZWEIFEL Chairman and Curator, Department of Herpetology American Museum of Natural History

BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY VOLUME 165 i ARTICLE 5 NEW YORK: 1980 BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY Volume 165, article 5, pages 387-434, figures 1-14, 1 table Issued September 22, 1980 Price: $3.10 a copy

This article completes Volume 165.

ISSN 0003-0090

Copyright © American Museum of Natural History 1980 CONTENTS Abstract .390 Introduction .390 Methods.391 Acknowledgments .391 The Geographic Setting .391 Previous Work in the Area .392 Systematic Accounts .393 Salientia.393 Bufonidae .393 Leptodactylidae .393 Hylidae .393 Ranidae .402 Microhylidae .403 Sauria .413 Gekkonidae .413 Pygopodidae.414 Agamidae.414 Scincidae.415 Varanidae,. 422 Biogeography.422 The Geologic Setting . 422 Faunal Distributional Relationships . 423 Endemism .423 Distributional Relationships within New Guinea . 423 Faunal Relationship to New Britain .428 History of the Peninsular Fauna . 429 Literature Cited .430 390 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 165

ABSTRACT The Huon Peninsula forms the eastern end of on the highest peaks, montane grassland expand- the Finisterre-Saruwaged mountainous region, ed and cooler temperatures prevailed. It is hy- nearly 300 km. long, which rises to 4000 m. above pothesized that these altered climatic conditions the sea and nearly that far above adjacent low- permitted immigration of such upland species as lands. Thirty-eight species of frogs and 52 of liz- are present and now, with changed climate, exist ards are recorded from the Peninsula, many of in disjunct populations. Climate did not change them for the first time. Endemism is slight: none sufficiently to permit immigration of high montane of the lizards and only three of the frog species species. Lowland species-especially skinks- appear to be endemic. Most of the lizards (92%) would have become established shortly after the and frogs (54% of non-endemic species) are lowland mass first appeared above the sea, and the land forms, although many of these range into more balanced lowland fauna would have been uplands as well. Populations of four species of achieved when the Ramu-Markham area became lizards and 16 of frogs evidently are disjunct from dry land. The scarcity of endemic forms suggests conspecific populations elsewhere, almost all of that the Finisterre-Saruwaged region has not been which are in mountains to the southwest of the isolated long enough or continuously enough to Peninsula, across the Ramu-Markham River Val- permit speciation of isolated populations. The ley. The disjunct species are ones with lower el- frog and lizard faunas provide no evidence of a evational limits at about 1000-1400 m., some low- former land connection between New Britain and er. Frog species restricted to high mountain the Huon Peninsula, and the scarcity of lowland forests or grasslands elsewhere in New Guinea New Guinea frog species in New Britain argues are absent from the Peninsula. against any such connection. Geological evidence indicates that the mountain One new species of microhylid frog-Cophix- mass of which the Huon Peninsula is part has alus pipilans-is described, and distributional, undergone 3000 m. of uplift since the late Plio- ecological, and systematic notes are given for cene, and that for part of its existence the area many of the species. was insular. In the Pleistocene, glaciers existed

INTRODUCTION The Seventh Archbold Expedition to New sen, 1978) and one report on vegetation (Cos- Guinea, led in 1964 by Hobart M. Van Deu- tin, Hoagland, and Lendon, 1977) have been sen, established collecting stations on the published. The present report derives from Huon Peninsula ranging from sea level to a manuscript submitted to Mr. Van Deusen 3500 m. For more than six months, from in 1972, now extensively revised to meet the April to October, members of the Expedition circumstances of its independent publication collected both botanical and zoological ma- and to incorporate information gathered terials, emphasizing among the latter mam- since then. mals, amphibians, and reptiles. In the course The purposes of this report are to record of a separate trip to New Guinea, I partici- the species of frogs and lizards known to oc- pated in the Expedition for a little more than cur on the Peninsula, to provide pertinent two weeks, June 17 to July 3. It was Mr. Van notes on ecology, systematics and distribu- Deusen's plan to organize a multi-authored tion, and to examine the frog and lizard fau- report on the Expedition, to which various nas in a biogeographic context. There have specialists would contribute chapters based been no comprehensive faunal reports on on their studies of the Expedition's collec- amphibians or reptiles from major regions of tions and, in some instances, their experi- New Guinea, such as are available for birds. ence as members of the Expedition. This The principal reason for this situation is that goal was frustrated by Mr. Van Deusen's even where good collections have been made death in 1976, and only a summary compiled (as on the several Archbold Expeditions, or from partly completed manuscript (Van Deu- by Fred Parker at numerous localities), there 1980 ZWEIFEL: HUON PENINSULA FROGS AND LIZARDS 391 remain so many unresolved systematic prob- Other fieldwork pertinent to the present lems that attempts at comparative faunal study took place in 1968 (supported by Na- studies would be woefully incomplete. It will tional Geographic Society grant no. 674) and be evident that this difficulty has encum- 1969 (as, a member of the R/V "Alpha Helix" bered the present study. Expedition to New Guinea, sponsored by the Scripps Institution of Oceanography, funded METHODS by the National Science Foundation). A series of species accounts, one for each Mr. Greenfield Sluder was my able field species of frog and lizard of which I have assistant in 1964. Members of the Alpha He- examined one or more specimens from the lix Expedition, notably Dr. Harold Cogger, study area, precedes the biogeographic dis- Dr. Herbert Dessauer, and Mr. Robert Sto- cussion. A few species were neither collect- rez, were co-workers in the field and con- ed by the Archbold Expedition nor examined tributed valuable specimens and much help. by me from other collections. I have not pre- Mr. John Womersley, then Chief of the Di- pared accounts of these, but all are included vision of Botany, Lae, was a never-failing in table 1 and some are treated in discus- source of logistical assistance. Data gathered sions. from specimens borrowed over many years Specimen numbers unaccompanied by for several projects from various museums museum abbreviations refer to specimens in contributed to this study. For their kindness the American Museum of Natural History. in making long-term loans, I am particularly Other museum names are abbreviated as fol- indebted to Mr. Michael Tyler, Dr. Ernest lows: BBM, Bernice P. Bishop Museum, E. Williams, and Dr. Alan Ziegler. Mr. Tyler Honolulu; BMNH, British Museum (Natural and Dr. Allen Greer provided helpful criti- History), London; MCZ, Museum of Com- cism of the manuscript. parative Zoology, Harvard University; MSNG, Museo Civico Storia Naturale, Gen- THE GEOGRAPHIC SETTING oa; SAM, South Australian Museums, Ade- The Huon Peninsula juts into the Solomon laide; YPM, Yale Peabody Museum, Yale Sea, forming the northern boundary of Huon University. Gulf and the southern boundary of Vitiaz Other abbreviations and notations used in- Strait. New Britain, across the strait, is only clude: TL (tibia length, from heel to outer 90 km. away at the closest point (fig. 1). The surface of knee); SV (length from snout to Peninsula is the eastern end of a rugged, vent); E-N (distance between anterior cor- mountainous region, the Finisterre and Sa- ner of eye and center of external naris); IN ruwaged ranges (elevations to 4100 m.) sep- (distance between centers of external nares); arated from the central ranges to the west by HW (head width, measured at level of tym- the low-lying Markham-Ramu River Valley panum); Eye (horizontal diameter of orbit, (maximum elevation, about 300 m.), and from anterior to posterior corner). The sym- from the Adelbert Mountains to the north by bol "±+" precedes one standard error of the the similarly low Gogol River drainage. The mean. limits of the study area were dictated by the area covered by the Expedition. From the ACKNOWLEDGMENTS standpoint of biogeography it would be more reasonable to study the Finisterre-Saru- My primary debt is to Mr. Hobart M. Van waged region as a whole, but in fact it makes Deusen, late Archbold Assistant Curator, for little difference to restrict the study to the his invitation to join the Seventh Archbold Huon Peninsula (defined by a north-south Expedition at its Gang Creek Camp on the line touching the western edge of Huon Huon Peninsula. My fieldwork in New Gulf), for scarcely anything is known of the Guinea in that year (1964) was supported by herpetofauna of the Finisterre Mountains, National Science Foundation grant GB-2217. west of the Huon Peninsula. 392 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 165

&3 14'0 150

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LOCALITIES ON THE HUON PENINSULA: the Huon Peninsula, and Mehely (1901) used Almost all the localities recorded in the fol- specimens from this region in an important lowing species accounts are mapped and in- revisionary work. Most herpetological col- cluded in the gazetteer in Van Deusen (1977). lections made in German New Guinea at this Exceptions are the following: Bukaua (40 time, however, were assembled to the km. E. Lae, sea level); Lialun (given as "bei west-around Astrolabe Bay and at sites Kap Konig Wilhelm" by Vogt [1911, p. 420], along the north coast and up the Sepik River. equivalent or close to Sialum); Sakimbang Following the change of government that (not found, = Selimbeng?); Tobo (not found, took place during the First World War, her- probably = Tobou); Tuwap (not found, prob- petological fieldwork languished, and it was ably = Tewep); Ulap (12 km. N. Kabwum); not until military activities in World War II Waran (8.5 km. S., 6.5 km. E. Kabwum). fortuitously brought herpetologists to the area that significant collecting resumed. PREVIOUS WORK IN THE AREA Loveridge (1948) summarized collections of New Guinea amphibians and reptiles in the Herpetological exploration of the Huon United States National Museum and the Mu- Peninsula dates from about the beginning of seum of Comparative Zoology, and included the present century, when persons working all specimens from the Huon Peninsula in at mission stations and administrative cen- these museums at that time. The most no- ters established during the period of German table collection was made by W. H. Stickel, rule brought collections to Europe. Lonn- but other persons contributed as well. berg (1900) and Vogt (1911) published an- Revisionary studies of varied scope (e.g., notated lists that included specimens from Brown, 1953; Brown and Parker, 1977; Par- 1980 ZWEIFEL: HUON PENINSULA FROGS AND LIZARDS 393 ker, 1934; Tyler, 1965, 1968; Zweifel, 1969, coastal fringe; the montane fauna was vir- 1972a, 1972c) have treated frogs and lizards tually unknown. It is not surprising, there- from the Huon Peninsula, but the fauna has fore, that the collections made in the interior been poorly known. Previous to the Seventh of the Peninsula by members of the Seventh Archbold Expedition, collectors had done Archbold Expedition substantially increase little more than sample the fauna of the our knowledge of the fauna.

SYSTEMATIC ACCOUNTS SALIENTIA adult individuals (the majority males) found BUFONIDAE the largest male to measure about 59-60 mm. snout-vent length, with the modal length Bufo marinus (Linnaeus) about 43 mm. (estimated from Tyler's fig. Lae (52591-52592+4, 66957-66969). 5).1 The AMNH specimen measures 61 mm., This introduced species is abundant in the and thus exceeds all the males measured by Botanic Garden in Lae. Zug et al. (1975) re- Tyler. Tyler reported that the male holotype ported marinus from Lae and some localities of Hyla papuensis Werner (from "German to the north of Lae, on the Huon Peninsula, New Guinea"), a name Tyler synonymizes but had no records for more inland and east- with amboinensis, measures 54 mm. Al- erly localities. though within the range of the Mabilabol sample, this size still is attained by no more than three or four individuals in that large LEPTODACTYLIDAE sample. There seems little doubt that am- Lechriodus aganoposis Zweifel boinensis is a much smaller frog in the Ma- bilabol region than to the east in the Huon Numbut, 1340 m. (75790, 75791); Gang Peninsula region. The question of the status Creek, 1340 m. (74646). ofpapuensis may need to be reopened when The specimens listed are the holotype and more eastern material becomes available. two of the paratypes (Zweifel, 1972c). The The call of L. amboinensis has not been species is known elsewhere from moderate described. My rather poor recording (fig. 2) elevations in the central mountainous chain gives a general impression of the call but is as far west as Sibil in the Star Mountains, of little use for fine analysis. The call lasts about 50 km. into Irian Jaya. Farther'west it about 2 seconds (in the example illustrated) evidently is replaced by a closely related and is comprised of 33 closely spaced notes species, Lechriodus platyceps Parker. about 0.05 sec. in length. The energy peak is at about 2500 Hz. The information may be HYLIDAE of use in verifying the synonymization of H. Litoria amboinensis (Horst) papuensis when calls of amboinensis from the south coast of New Guinea become avail- Busu River, ca. 24 km. (by road) N. Lae, able. However, a highland member of the 200-250 m. (80902). same (peronii) species group (Tyler and Da- This frog was calling at night from high in vies, 1978), Litoria darlingtoni (Loveridge), a banana plant overhanging a still pool in a has a chattering call much like that of the small valley; others called nearby. Tyler (1968) recorded the species from Sattelberg 1 There is some confusion about the maximum length. on the Huon Peninsula. The histogram shows males as large as 59-60 mm. (data Tyler (1968) studied a large number of am- were pooled in 2 mm. increments), but Tyler and Davies boinensis from Mabilabol in the southern (1978) gave maximum length of the male for the peronii drainage of Irian Jaya, and in a sample of 341 group as 54 mm. 394 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 165

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---- o r- I I_ I .2 .4 .6 .8 1.0 1.2 1.4 1.6 1.8 2.0 TIME IN SECONDS FIG. 2. Audiospectrogram (50 Hz filter) of call of Litoria amboinensis, recorded Aug. 25, 1968, 24 km. N. Lae, Morobe Prov., Papua New Guinea; AMNH 80902, temperature not recorded, Dept. Her- petology tape no. 187. amboinensis I recorded. It may be that calls Litoria angiana (Boulenger) in allopatric species of this group are not well Mountains at head of Kua differentiated. River Valley, 1610-2160 m. (75837); Avengu (75838-75840); In their diagnosis and definition of the pe- Indagen, ca. 1830 m. (75841, 75842); vic. Pin- ronii species group, Tyler and Davies (1978, diu, 1320 m. (BBM 2922); Kwama River, 20 p. 39) stated: "Flash markings of black and mi. SW. Kabwum, 2000 m. (BBM 5208). yellow or orange are present in all individu- These specimens agree well with the def- als," and the "posterior surface of the thighs inition and description given by Tyler (1968, ... are black and yellow or black and or- pp. 33-38). The species ranges along the cen- ange." This is not true of my specimen of tral mountainous spine of New Guinea from amboinensis, nor was it true of amboinensis from the Vogelkop Peninsula of Irian Jaya the Vogelkop Peninsula to the vicinity of described by Brongersma (1953): "Posterior Garaina (Tyler, 1968; Zweifel, 1976). The present specimens are the first recorded from surfaces of thigh and tibia uniformly greyish the Huon Peninsula. Because this species is except close to the vent." associated with montane streams, the popu- Tyler and Davies (1978) did not mention lation of the Huon Peninsula presumably is a distinctive and presumably synapomorphic character of the peronii group-the shape of disjunct from the main body of the species. the pupil. In living amboinensis, darlingto- Litoria eucnemis (Lonnberg) nii, and rothi that I have examined, the pupil is diamond-shaped when contracted, in con- Busu River, 13 km. N. Lae (74702-74718); trast to the horizontal slit of all other Litoria ca. 24 km. N. Lae, 200 m. (80772). I have seen and the vertical slit of Nycti- The type locality, Sattelberg, is the only mystes. The pupil appears diamond-shaped other locality for this species on the Huon in a preserved L. everetti and evidently has Peninsula. Tyler (1968) provided specific lo- the same shape in the remaining species, pe- cality records, and Menzies (1976) referred ronii, for Barker and Grigg (1977) remarked to "scattered records . .. through the north- on its being the "shape of a cross." ern and southern lowlands. Because of the 1980 ZWEIFEL: HUON PENINSULA FROGS AND LIZARDS 395 possibility of confusion between this species statistics for seven adult male genimaculata and L. genimaculata, some discussion is re- from Morobe Province are: mean, 38.9 + quired. 0.55, range 37-40. Overlap in the samples is Menzies (1976) stated that eucnemis and slight, for only one eucnemis measures less genimaculata are "'not easy to distinguish" than 41 mm. and aside from noting that males of eucnemis Tyler (1968) characterized eucnemis as reach a body length of 48 mm. compared having fully webbed fingers. Few of my male with 41 mm. in genimaculata, he offers no specimens could be so characterized, as the distinguishing physical characteristics. Men- webbing in most does not reach to the disc zies describes the call of genimaculata as of the third finger; I would call them three- "a very quiet ticking, about 15 ticks in a se- quarters webbed. However, a large female ries lasting 2 or 3 seconds," whereas euc- (SV 61 mm.)-the specimen from Oomsis nemis gives a "rather louder . . . series of Creek examined by Tyler (AMNH 66964)- low-pitched notes at intervals sounding like has fully webbed fingers. The specimens of "waa-waa-waa." genimaculata have much less webbing, com- Tyler (1968) did not make diagnostic commonly about one-third webbed. The differ- parisons between eucnemis and genimacu- ences between the two samples are evident lata. He compared eucnemis only with enough when specimens can directly be com- species having fully webbed fingers, and gen- pared, although isolated specimens in the in- imaculata with "lowland species whose termediate size range might give pause. adults are within the size range of adult H. The single eucnemis I tape-recorded (oth- genimaculata." Evidently there was some ers were heard giving the same call) pro- confusion regarding the range of elevations duced groups of low, soft, chuckling notes occupied by eucnemis, for Tyler excluded it and finished the series with several louder, from his key to lowland species ("0-3.500 ft shorter, pulsed calls (figs. 3B, 3C). These ter- above sea level"), where genimaculata keys minal calls have the appearance of shorter, out, and included it only in his key to sub- more rapidly pulsed versions of the preced- montane species ("3.000-5.000 ft above sea ing groups. I suspect that these terminal level"). One of the localities Tyler cites for notes are the same as the "waa-waa-waa" eucnemis (Oomsis Creek, Morobe Province, calls Menzies described. Papua New Guinea) is at only 100 m. ele- A genimaculata that I recorded (again, vation. others were heard giving the same call) pro- If I have identified my specimens correct- duced soft, ticking notes (as described by ly, eucnemis and genimaculata are highly Menzies) at a rate of about 6 per second (fig. similar. Their color patterns are variable but 3A). The number of notes within six call much alike (see color photos in Menzies, groups varied widely-from nine to 65. The 1976, and Zweifel, 1977),2 and body propor- call is exceedingly soft; I had to place my tions commonly of use in distinguishing microphone within 10 cm. of the frog in order species of Litoria are of little help. My seg- to get a signal loud enough to be heard over regation is based on size and finger webbing, the other night noises. The softness of the bolstered by some information on mating calls of the two species is no doubt related calls. to the absence of a vocal sac (confirmed by Among 36 adult male eucnemis (all have me), an unusual condition in the genus Li- thumb pads) from Madang and Morobe prov- toria (Tyler, 1971). inces, the range in snout to vent length is 38 The specimen of genimaculata that I re- to 51 mm., mean 46.7 + 0.45. Comparable corded has a snout-vent length of 37 mm., the eucnemis, 49 mm. Finger webbings con-

2 form to the norms discussed above. I con- The frog illustrated in color in Zweifel, 1977, p. 29, as "Litoria genimacotata" is L. eucnemis from Sempi, sider that the information on vocalizations, Madang Province. although meager, validates the species seg- 8 A 1$ . P .0 titi . . *44¶ih ¾ 5¾ 6 t

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.2 .4 .6 .8 1.0 1.2 1.4 1.6 1.8 2.0 TIME IN SECONDS FIG. 3. Audiospectrograms (300 Hz filter) of calls of Litoria. A. L. genimaculata, recorded August 15, 1964, 21 km. NW. Lae, Morobe Prov., Papua New Guinea; AMNH 74725, air 24.50 C., Dept. Herpetology tape no. 143. B. L. eucnemis, recorded Aug. 4, 1969 at Wanuma, Adelbert Mountains, Madang Prov., Papua New Guinea; AMNH 82611, air 21.20 C., Dept. Herpetology tape no. 189. C. Same call as B, but terminal notes of series. 1980 ZWEIFEL: HUON PENINSULA FROGS AND LIZARDS 397

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0 .2 .4 .6 .8 1.0 1.2 1.4 1.6 1.8 2.0 TIME IN SECONDS FIG. 4. Audiospectrogram (50 Hz filter) of call of Litoria infrafrenata, recorded Aug. 3, 1964, at Lae, Morobe Prov., Papua New Guinea; no specimen, air 240 C., AMNH Dept. Herpetology tape no. 142. regation on the basis of size and finger web- (Peria Creek, Tyler, 1968). Elsewhere, gen- bing. imaculata evidently ranges over the south- Litoria genimaculata and L. eucnemis ap- ern lowlands of New Guinea (Menzies, pear to be sibling species with broadly over- 1976). lapping distributions and at least some mi- crosympatry. I have collected them together Litoria infrafrenata infrafrenata at two places, and in neither instance did I (Gunther) distinguish the species in the field. I found Finschhafen (75956); Lae (52579, four genimaculata and three eucnemis in the 52580+ 19, 52587, 58690-58693, 74729- Markham Valley, 21 km. northwest of Lae, 74731). elevation lower than 75 m. They were on low This common lowland species was not en- shrubs in forest at night beside a shallow, countered at the inland sites. One of the sand-bottomed stream no more than 5 feet specimens, a female, from Lae (74729) with wide. The stream vanished into the sand a snout to vent length of 141 mm., is the within a few yards of where the frogs were. largest L. infrafrenata I have collected and The other area of sympatry was similar, may be the largest specimen of a hylid frog. where two individuals were on bushes over Barker and Grigg (1977) accurately de- a stream about 24 km. north of Lae, eleva- scribed the call of L. infrafrenata as a "loud, tion about 200 m. harsh double note." The call is illustrated in figure 4. As do frogs of many species, L. in- Litoria genimaculata (Horst) frafrenata sometimes gives a "fright scream" Ca. 24 km. N. Lae, 200 m. (80773). when grasped (fig. 5). For discussion of this species, see the preceding account of L. eucnemis. This record and others for the lower Markham River Val- Litoria micromembrana (Tyler) ley within 16-24 km. of Lae are the north- Gang Creek, 1310-1370 m. (75898-75947, ernmost for the species on the north coast of 74668-74670, 99383-99424); Kotkin, 1220 m. New Guinea, separated by about 440 km. (75955); Selimbeng, 1370 m. (75948-75954). from the other published north coast record These frogs are readily identifiable as mi- on the mainland west of Goodenough Island cromembrana as defined by Tyler (1968, p. 398 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 165

8' Duellman (1967) used one of the Expedi- tion specimens (74668, "H. becki") in a study of chromosomes. 6' Litoria thesaurensis (Peters) N Pindiu, ca. 850 m. (75957, 75958); Gusiko I 4' (52576); Lae (66960, 74752, 80901). This widespread species is characteristic of coastal lowlands. See Menzies and Zug, 2 1979. Litoria wollastoni (Boulenger) .2 .4 .6 .8 1.0 Gang Creek, 1310-1370 m. (74662-74667, TIME IN SECONDS 75849-75897+18); Numbut, 1340 m. (75845); Mt. Rawlinson, 1830 m. (75846); north slope FIG. 5. Audiospectrogram (50 Hz filter) of of Mt. Rawlinson (75847); ridge above Gang fright screaim of Litoria infrafrenata, recorded Creek (75848); Pindiu, ca. 850 m. (75843, Aug. 3, 1964, at Lae, Morobe Prov., Papua New 75844). Guinea; At4NH 74730, air 240 C, Dept. Herpe- Litoria arfakiana of earlier authors (e.g., tology tape no. 142. Tyler, 1968, p. 39) includes two species that are so similar morphologically that they cannot be distinguished with certainty except by 131), excelpt that they evidently represent a... means of their distinctly different calls (Men- population of somewhat smaller body size. zies and Zweifel, 1974). The call that I tape- The range length from snout to vent of9 recorded at Gang Creek is that of the species adult malesin 28-33 (me 1 9mm.) for which Menzies and Zweifel (1974) use the compared sish a mm.gof3. 30.1toto 3 name wollastoni Boulenger. Hence, I refer with. s given by T ragersmaller smm all specimens to this species, although it is ilarity to yLetrTis sizeylens at possible that Litoria arfakiana (Peters and Liomtori mod (er),sutagrehee specimens Doria) also is represented in the series. with micro frombrthe Hond Piffeninul m Duellman's (1967) "H. arfakiana" is one ica) in the memanave (andrdiferfye,longero mod,t of and more crelatvel lanrge tra,lo. the Expedition specimens (no. 74666). Litoria wollastoni is notable for the con- All indiNviduals of thus speclies. color and pattern lo- tured were foindualstfound at nightthistspeciesoon bushesthtrIcap-or trees siderable variation in in beside or cal populations. I recorded the following theals "a one occasion I characterized onevariationsevening:among21 30frogsspecimensuniformcollectedgreenishin peep repeated sometimes several times, brown dorsally; four brown with a darker and on anc)ther as "a series of short, rasping tral chirps" (fit eldnotes). centa pattern on the back; three brown with Tyler's l 1968) re d f t the top of the head bright green from a mid- the ocular line to the tip of the snout and the of this spe-ciescies8)rangeraneodfromforothe SnowdisnowbMoun-Moun- back of the tarsus green; two all green except tains of eatstern Irian Jaya to Okapa in the for black canthal and supratympanic stripes. Eastern Hlighlands, and specimens studied The frogs call at night from bushes or trees by Dessauier, Gartside, and Zweifel (1977) A. & & & above turbulent streams. extend the range 2zu1% '1%.1Ikm. to tne1- soutnwest1- to Saureli in Morobe Province. The present specimens come from about 180 km. east of Nyctimystes disrupta Tyler Okapa andl 150 km. north of Saureli, and are Gang Creek, 1310-1370 m. (74818, 74819, the first recorded from the Huon Peninsula. 74824, 75991-75993); near Zangaren, 1370 m. 1980 ZWEIFEL: HUON PENINSULA FROGS AND LIZARDS 399

(75994-75999); Numbut (76000-76002); Kot- region (Tyler, 1963c; Bulmer and Tyler, kin, 1220 m. (76003). 1968; Woodruff, 1972), but the species is Tyler (1963b, pp. 111-112) showed that the widely distributed in the mountains of New syntypes of Nyctimystes papua (Boulenger) Guinea as attested by collections made by are not all conspecific and later (1963c, pp. the Sixth Archbold Expedition, by Fred Par- 118-120) he described N. disrupta, a ker, and by me. The range extends at least "species with a close affinity to N. papua," from Ialibu in the Southern Highlands Dis- from the Schrader Mountains and the Baiyer trict (elev. 1920 m.) to the vicinity of Wau River region. (elev. 1400 m.), Morobe District (Zweifel, The specimens from the Huon Peninsula 1976), and to mountains just south of the resemble disrupta more than papua, partic- mouth of the Markham River (Wagau, Her- ularly in the amount of finger webbing, a zog Mountains, 1100 m., AMNH 74771- character to which Tyler attributes signifi- 74779), about 65 km. from localities on the cance. Identification of the Huon Peninsula Huon Peninsula. Presumably the populations specimens as disrupta is tentative, pending of N. foricula and other Nyctimystes on the further studies on N. papua and its relatives. Huon Peninsula are disjunct from the popu- I recorded the following colors in life for lations south and west of the Markham and number 74818: dark green dorsally with red- Ramu river valleys, for these valleys appear dish brown flecks and other markings; dorsal to offer no suitable habitat for frogs that surfaces of hind limbs dark green, mottled breed in fast-flowing streams. and banded with reddish brown; chin and Specimens that I captured were calling chest pale green with reddish brown mark- from bushes and shrubs by small streams. ings; abdomen, groin, anterior and posterior An individual from Gang Creek (74767) had surfaces of thighs, and webbing of hands and the following colors in life: dorsal body sur- feet purplish red; iris green; palpebral vena- face bright green with some green and yel- tion golden but faint. The holotype (in pre- lowish green mottling; anterior and posterior servative) had "copper" ventral surfaces to surfaces of thighs immaculate cadmium yel- the limbs, hands and feet, and "lilac" ab- low; undersurfaces of hind limbs less bright domen (Tyler, 1963c, p. 119), which suggests yellow; ventral body surfaces lemon-yellow; a coloration in life similar to that of the spec- upper arm green, forearm yellow above and imens from the Huon Peninsula. below; iris virtually black, pupil indistin- This species has not previously been re- guishable; palpebral venation composed of corded east of the Western Highlands Dis- oblique, parallel, golden lines. trict, although it is, in fact, fairly widely dis- Duellman (1967) published chromosome tributed. I have collected specimens referable numbers determined from Expedition speci- to disrupta at about 13 km. southwest of Go- mens (Nos. 74767, 74769). roka, 2190 m., Eastern Highlands Prov., and at Wagau, 1100 m. (about 24 km. southwest Nyctimystes kubori Zweifel of Lae), Morobe Prov. Duellman (1967) reported chromosome Gang Creek, 1310-1370 m. (74794-74801, numbers from Expedition specimens ("N. 75970-75984, 75986-75990, 100861-100869); papua" Nos. 74819, 74824). Zangaren, 1370 m. (75985); Tumnang (74793). These specimens are the first of their Nyctimystes species recorded from the Huon Peninsula. foricula Tyler Most records published previously are for Gang Creek, 1310-1370 m. (74767-74770, areas far to the west (mountains flanking the 75964-75967, 75969); Numbut, 1220 m. Wahgi Valley and the Schrader Mountains: (75962); Kotkin (75963); Tumnang, 1340 m. Tyler, 1963b, p. 109; Tyler, 1963c, p. 123; (74764-74766). Zweifel, 1958, p. 21), but there now are spec- Most literature records for this species re- imens in several museums from localities fer to localities in the Schrader Mountains closer to the Huon Peninsula. I have, for ex- 400 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 165 ample, collected kubori in the vicinity of that it is a species distinct from pulchra. I Wau (Zweifel, 1976) and at Wagau, 1100 m. have collected the true semipalmata only at (AMNH 74780-74792). The latter locality is Kokoda (type locality) and at Garaina, about in the Herzog Mountains 65 km. from Gang 130 km. northwest of Kokoda (Zweifel, Creek, across the Markham River Valley. 1976). Nyctimystes semipalmata is morpho- I found N. kubori calling from riparian logically similar to N. cheesmani, with which vegetation in company with N. foricula, Li- it is sympatric at Kokoda and Garaina. In toria wollastoni, and L. micromembrana. life, they are readily distinguished by differ- In life, different individuals of kubori from ences in calls and pigmentation (webbing the Huon Peninsula showed a variety of dor- bright orange in semipalmata, gray in chees- sal ground colors, ranging from light yellow- mani). ish brown to gray and dark brown. The dor- sum was immaculate, lightly spotted with Nyctimystes sp. darker gray or brown, or rather heavily mottled. The only bright coloration was in the Pindiu (75959); ca. 24 km. N. Lae, 250 m. webbing of the feet, which in some individ- (80933-80939). uals was largely orange, although in others These specimens possibly represent an un- only a peach tint relieved the otherwise gray described species of the cheesmani3 group webbing. The vertical pupil could scarcely (Menzies, 1976). In my now somewhat out- be distinguished against the blackish brown dated key to Nyctimystes (Zweifel, 1958), iris. The densely reticulate palpebral vena- the specimens would be identified as N. tion was grayish gold. montana Parker (= N. cheesmani Tyler, a Duellman (1967) used Expedition speci- substitute name). I have collected what I mens in determining chromosome numbers think is true cheesmani at several localities: (Nos. 74794, 74800, 74801). Garaina and the vicinity of Wau (Morobe Province); Kokoda (Northern Province); and Nyctimystes pulchra Baiyer River Wildlife Sanctuary (Western (Wandolleck) Highlands Province). Specimens from these Mt. Rawlinson, between Maran and Zan- localities invariably had a dark brown iris, so garen (75961); near Zangaren, 1370 m. dark that the pupil was scarcely visible. In (76004); Pindiu, 850 m. (76005). contrast, the iris of the Pindiu specimen (as Tyler (1964b) resurrected the name Hyla seen in a color photograph) was russet, and pulchra Wandolleck and showed that on the in the Lae specimens (my field notes) was basis of the original description, the species gray-brown with a golden tint, the pupil quite belonged in Nyctimystes rather than Hyla. distinct. He was unable to carry his investigation fur- The call of N. cheesmani is a monoto- ther, however, because the types had been nously repeated series of quacking notes (fig. destroyed and no additional specimens had 6B). The call of the pale-eyed form is similar been collected in the region of the type lo- (fig. 6A), but in my single recorded sample cality (Torricelli Mountains). seemed to be given with less regularity. In 1966 Dr. Jared Diamond collected a Also, individual notes are longer and less large series of these frogs (AMNH 77857- rapidly pulsed. In the examples illustrated, 77875+56) in the Torricelli Mountains, and the pulse rate in N. cheesmani is about 100 comparison of these specimens with others per sec., compared with 80 per sec. in the leaves me no doubt that the specimens I for- pale-eyed form. Differences cannot be attrib- merly referred to Nyctimystes semipalmata Parker (Zweifel, 1958, pp. 38-42) are the same species as Wandolleck'spulchra. Also, I Menzies (1976) emended this name to cheesmanae. since writing my 1958 paper, I have exam- This was an unjustified emendation, since the XVIth Con- ined the holotype of semipalmata and have gress of Zoology in 1963 reduced Article 31 of the Inter- collected topotypes, and I do not question national Code to the status of a Recommendation. 1980 ZWEIFEL: HUON PENINSULA FROGS AND LIZARDS 401

~t4 W. p V

0 .2 .4 .6 .8 1.0 1.2 1.4 1.6 1.8 2.0

I If F I I_ B

.2 .4 .6 .8 1.0 1.2 1.4 1.6 1.8 2.0 TIME IN SECONDS FIG. 6. Audiospectrograms (300 Hz filter) of calls of Nyctimystes. A. N. sp., recorded Aug. 25, 1968, 24 km. N. Lae, Morobe Prov., Papua New Guinea; AMNH 80933, body 22.80 C., Dept. Herpe- tology tape no. 187. B. N. cheesmani, recorded July 7, 1968, at the Baiyer River Wildlife Sanctuary, Western Highlands Prov., Papua New Guinea; AMNH 8094 1-80945 vouchers (individual recorded not determined), air 200 C., Dept. Herpetology tape no. 182. uted to temperature, as the calls of the cooler Province, and have seen color photographs frog were more rapidly pulsed. of specimens much like it from Kiunga, I refrain from naming the possibly new Western Province (MCZ field no. 30605) and species because the identity ofN. cheesmani Ramu Gorge (Province?) (MCZ field no. needs to be confirmed through tape-record- 22138), collected and photographed by Fred ings of the calls of topotypes, and the status Parker. Menzies (1976, pl. 9a) illustrated of other named forms similarly needs to be what may be the same species. Seven spec- clarified. I have collected what I think is the imens in the South Australian Museum undescribed species near Saureli, Morobe (SAM 4250) from Finschhafen may represent 402 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 165

N I4

.... 2 t.

0 Jlg .2 .4 .6 .8 1.0 1.2 1.4 1.6 1.8 2.0 TIME IN SECONDS FIG. 7. Audiospectrogram (50 Hz filter) of call of Rana daemeli, recorded July 17, 1964, at Lae, Morobe Prov., Papua New Guinea; AMNH 74864, air 25.50 C., Dept. Herpetology tape no. 140. this species, but without data on eye color jimiensis Tyler. None of these has been in life their status cannot be confirmed. found within the area covered by this report. The several remaining species, which may RANIDAE informally be called the Rana papua group, are similar enough so that associating the Platymantis papuensis Meyer many names in the literature with animals in Masba Creek, 610 m. (75792-75803); Gu- hand or observed in the field is difficult or siko (52574); Lae (52575, 52581, 52584, impossible. 52586, 66961, 66962, 74828-74832, 75804, The type locality of Rana daemeli is Cape 102961-102973). York, Queensland, Australia. Recent au- This species is widespread at low eleva- thors (Cogger, 1975; Barker and Grigg, 1977) tions throughout northern New Guinea have recognized only this one species of (Zweifel, 1969), and was absent from all but Australian Rana, although Loveridge (1948, the lowest of the inland camps on the Huon p. 413) considered Rana grisea to be there Peninsula. The binomial is used because of as well. Limited evidence indicates that evidence that supposed subspecies of pa- Rana daemeli ranges through the lowlands puensis in the Bismarck Archipelago and of southern New Guinea and along the north Solomon Islands are actually one or more coast from Milne Bay to Finschhafen on the distinct species (Menzies, unpublished data). Huon Peninsula (Menzies, 1976, p. 25). Men- zies describes the call as "rather like the Rana daemeli (Steindachner) noise made by a duck," and Barker and Grigg describe a "low, reedy quacking." Lae (74863, 74864, 81292, 81293). The call of the frogs I identify as daemeli The systematics of New Guinea Rana are (fig. 7) fits these descriptions. I have record- in a confused state. Only one species-Rana ed it at Wipim, Western Province, directly grunniens Daudin-is sufficiently distinctive north of Cape York, as well as at Lae. I did morphologically that it can be recognized at not hear the species in the vicinity of Alex- a glance. The widespread Rana arfaki Meyer could be confused with the little-known R. ishafen, Madang Province, nor at Maprik, 1980 ZWEIFEL: HUON PENINSULA FROGS AND LIZARDS 403

East Sepik Province, which accords with the sula specimens come from both higher and distribution Menzies postulates. lower elevations. In this regard, it is perti- Tyler (1972) considered Rana daemeli a nent to note that the type-locality of R. gri- synonym of Rana papua Lesson. The type sea is given as "Went-Gebirge, + 1300 m." locality of papua, Waigeo Island, is off the (Van Kampen, 1913). I do not question Men- northeastern tip of New Guinea, far from the zies's conclusion that highland and lowland known range of daemeli. Despite frequent forms are different species (based on vocal- use of the name papua in the literature, it izations), but assignment of the name grisea probably will not be possible to determine to the high-elevation species may be incor- which (if any) of the mainland species is pa- rect. As is the case with Rana papua, the pua until fresh specimens are collected and problem is not likely to be adequately re- recordings of the call made at the type lo- solved unless topotypes can be collected and cality. tape-recorded. Duellman (1967) reported the chromosome Rana grisea Van Kampen number of one of the Huon specimens Masba Creek, 610 m. (75815-75820); Pin- (74850). diu, 790-880 m. (75806-75814); Zangaren, Rana sp. 1370 m. (75822); Tumnang, 1340 m. (74849); Numbut, Mt. Rawlinson, 1340 m. (74850, Pindiu, ca. 880 m. (75805). 75821); Gang Creek, Mt. Rawlinson, 1310 m. I identify this specimen with the "pond- (74861, 74862); Indagen, ca. 1830 m. (75823- dwelling, lowland" species that Menzies 75834); Waran, ca. 1890 m. (75835, 75836). (1976, pp. 25-26) reports sympatric with The name Rana grisea has been used for Rana daemeli and the lowland, stream- frogs from high elevations throughout much dwelling species (similar to grisea, see of New Guinea. These are large, long-legged above) in the Botanic Garden at Lae. The frogs readily distinguished from R. daemeli chief distinguishing characteristic is the call, and Rana sp. (see following species account) which he describes as "single notes rather in size, proportions, and some aspects of col- like plucking a flat string on a violin." I offer or pattern. However, frogs closely similar in here an audiospectrogram (fig. 8) of the call size, proportions, and pattern also occur at of this species, recorded at Wau, Morobe low elevations over much of New Guinea. Province. Menzies (1976, pp. 34-35) characterized this This species ranges along the north coast lowland form as a stream-breeding species of New Guinea at least from Popondetta, that calls with "a loud, rattling series of Northern Province (Menzies) to Maprik, notes usually lasting two to three seconds." East Sepik Province (my specimens and Apart from differences in vocalization, Men- tape-recording). Menzies regards this species zies finds nothing to distinguish the lowland and daemelii as indistinguishable morpholog- form from grisea except for yellow ventral ically except for the occasional presence in coloration in young grisea, lacking in young the north coast species of a yellow vertebral of the lowland form and in adults of both line. I find other differences in pigmentation species. and proportions that, at least in my samples Menzies mentions that the lowland form from Lae and the lower Markham Valley, occurs at Lae (where I have heard but have permit segregation. not collected it), in sympatry with R. daemeli and Rana sp. (see following account). The MICROHYLIDAE identity of the specimens I record here as grisea is questionable. Menzies states that Cophixalus biroi (Mehely) grisea "probably does not occur below 1400 Gang Creek, 1370 m. (74904); Sakimbang, m.," whereas the lowland form is found 1390 m. (84504). "perhaps up to 1400 m." The Huon Penin- Information on this species is summarized 404 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 165

8 AMNH 83000 and 83001, collected at the type locality on July 13 and 14, 1969; AMNH 83003, Wanuma, Adelbert Mountains, 700 6 mI., Madang Province, August 6, 1969; AMNH 83005-83008, 3.5 km. northwest of Alexishafen, elevation 40 m., Madang Prov- N 4 ince, September 14 and 15, 1969; MCZ 89228-89234, 4 km. north of Lae, Morobe Province, collected by Fred Parker, March 2 ;:. 3, 1974. DIAGNOSIS: A small species of Cophix- I -. alus (males to 18.5 mm. snout-vent length, oZ. females. . . .to-l22.3,- mm.)* .- that- differs from most .2 .4 .6 .8 1 species of the genus in having the first finger short, less than half the length of the second, TIME IN SECONDS and without a broadened disc (fig. 9), where- FIG. 8. Audiospectrogram (50 Hz filter) of call as the remaining fingers have well-developed of Rana sp., recorded July 22, 1968, at Wau, Mo- discs smaller than those on the toes. Two rotbe Prov., Papua New Guinea; AMNH 81291, species share the diagnostic character of the sulbstratum (lily pad) 19.40 C., Dept. Herpetology first finger (fig. 9) with C. pipilans: C. ateles tap (Boulenger) and C. shellyi Zweifel. The new species differs from the latter in leg leingth (TL/SV 0.53 or greater inpipilans, maximum in Zweifel (1979b). The type locality is Sat- of 0.53 in shellyi) and in having more widely telIberg, on the Huon Peninsula. The species spaced nostrils (E-N/IN minimum 0.76 in occurs in the north coast ranges of Papua pipilans, maximum 0.77 in shellyi). Cophix- ew Guinea, westward into Irian Jaya. alus ateles resembles C. pipilans in its E-N/ IN ratio, although ateles averages and ranges Cophixalus cheesmanae Parker higher, but the leg length of ateles is mark- shorter: maximum TL/SV in Lae (52589); ca. 24 km. N. Lae (81075(81075,0.49,edly minimum in pipi/ans, 0.53. Theate/es,three 811076); Finschhafen (SAM 4249, 16 speci- 0f iiu npipln,05 h he 076); species are at greater in a mc >ns). section that comparedfollows. length This is a common species of foothill rain folrest, where males call from low shrubs at Menzies (1976) illustrated the species male,val callingc isnowheneiencaptured;externally.the sub'gular nitght. ,,. , ,,,x ~~~~~vocal sac iS not evident externally. in color, and Zweifel and Parker (1977) pre- Head moderately wide (HW/SV 0.36) se]nted an audiospectrogram of the call. For with s oblique, nea flat orea36e- diXagnostic and distributional information, n, obtusely obtusely rounded se e Zweifel (1979b). ' thusgion, rostralis, andpointednostrilssnout,much nearercan-to snout tip than to eye; internarial distance Copixaupiplas,ewspeie wider than distance from eye to naris (E-N/ HOLOTYPE: AMNH 83004 (field no. RZ IN, 0.823; IN/SV, 0.106, E-N/SV, 0.087; 8724), an adult male, collected by R. G. eyes moderately large (Eye/SV, 0.124); eye-

Zvveifel (in company with R. Storez) in the lid one-half width of interorbital span; tym- Vic:iinity of Sempi, 9 km. north of Alexishaf- panum obscure, about one-half diameter of en[, Madang Province, Papua New Guinea, eye. elevation between sea level and 50 m., on Dorsal surfaces of body and legs smooth- At igust 25, 1969. no obvious skin folds or warts. PARATYPES: YPM 5281-5283, collected Fingers unwebbed, relative lengths at the same time and place as the holotype; 3>4>2>1, the first finger less than one-half 1980 ZWEIFEL: HUON PENINSULA FROGS AND LIZARDS 405

A B C

FiG. 9. Palmar views of right hands of Cophixalus (scale lines marked in mm.). A. C. pipilans, AMNH 83004 (holotype); B. C. shellyi, AMNH 58551 (holotype); C. C. ateles, British Museum (Nat. Hist.) 1947.2.12.7 (paralectotype). the length of the second; second, third, and low eye-dark; a short, dark mark from pos- fourth fingers with enlarged discs with ter- terior corner of eye above tympanum, and a minal grooves; first finger tip rounded with- similar dark mark at same level on body just out disc or groove; disc of third finger slight- posterior to arm insertion; a dark mark at the ly less than twice the width of the penultimate wrist; legs with no distinct darker markings phalanx, but larger than other discs; low, but anterior and posterior surfaces of thighs rounded subarticular and inner metacarpal obscurely mottled; chin and chest with a tubercles barely indicated. Toes without dark stipple of melanophores broken by webbing or fringes, relative lengths small clear spots; abdomen largely pale; un- 4>3>5>2>1, third much longer than fifth; dersides of hind legs light brown with pale toe discs larger than those on fingers, that of mottling. the fourth toe about twice the width of the MEASUREMENTS (IN MM.): SV, 16.1; TL, penultimate phalanx; low, rounded subartic- 9.1; HW, 5.8; Eye, 2.0; E-N, 1.4; IN, 1.7; ular tubercles; inner metatarsal tubercle disc of third finger, 0.6 (penultimate phalanx, barely indicated. Hind legs long (TL/SV, 0.35); disc of fourth toe, 0.8 (0.4). 0.565). VARIATION IN THE TYPE SERIES: The Maxillae eleutherognathine (not meeting in largest male in the series measures 18.5 mm. front of premaxillae); no vomerine, premax- SV, and males are mature (found calling) at illary or maxillary teeth; sternum cartilagi- as least as small as 16.1 mm. Six gravid fe- nous; clavicles, procoracoids and omoster- males measure between 18.1 and 22.3 mm. num absent. SV. The principal diagnostic proportions Ground color of dorsal surfaces of body vary as follows: TL/SV, mean 0.563 + 0.006 and limbs brown in preservative, paler with (0.53-0.62), N = 18; E-N/IN, mean 0.806 + a dark stipple of melanophores on the side 0.009 (0.72-0.87), N = 18. A comparison of of the body; an indistinct, dark hourglass- the subsamples from Morobe and Madang shaped mark on the back of the head and provinces shows no significant differences in shoulder region; side of head-snout to be- these proportions. The first finger is typically 406 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 1 65

A (18) B (6)

C(8)

D (23)

E (5) I1

.40 .45 .50 .55 .60 TL/SV

A(18)

B (6)

C (8)

D (23)

E(5) I fL ,

I I I I I I I I I I I I I I I I,I I I I I I I I .50 .60 .70 .80 .90 EN/IN FIG. 10. Ratios of tibia length to snout-vent length and of eye-naris distance to internarial distance in five samples of Cophixalus. Ranges indicated by horizontal lines, means by vertical lines, with boxes marking two standard errors on each side of mean, sample sizes in parentheses. A. C. pipilans, entire sample; B. C. shellyi, Huon Peninsula; C. C. shellyi, vicinity of Wau; D. C. shellyi, Highlands region; E. C. ateles, type series.

short in all specimens. However, in some the COMPARISONS WITH OTHER SPECIES: Co- tip is disclike (although not perceptibly phixalus pipilans is so closely similar to C. broadened) and bears a faint terminal shellyi in size, color, and nature of the first groove. finger that when I encounteredpipilans in the The dorsal color is brown to yellowish tan field I took it for shellyi, though puzzled by in life, the face mask black. When frogs are the appearance virtually at sea level of a in light phase (as at night), the intensity and species known before from elevations of contrast of the face mask are reduced. One from 1200 to 2700 m. The almost complete specimen has the middle of the back much separation between pipilans and shellyi in paler than, and sharply distinguished from, certain proportions (fig. 10), and evidence for the sides. The groin and anterior and poste- distinctive vocalizations (following section) rior surfaces of the thighs have a pink to red- lead me to regard the two as different dish orange color. The throat and chest are species. No instances of sympatry are gray with pale flecks. known, but typical examples have been tak- 1980 ZWEIFEL: HUON PENINSULA FROGS AND LIZARDS 407

N I 4.

.- -- mllr. 0 ---Ir. _I I u = u ,-u-. . *I I 0 -- .2 .4 .6 .8 1.0 1.2 1.4 1.6 1.8 2.0 TIME IN SECONDS FIG. 11. Audiospectrogram (50 Hz filter) of call of Cophixalus pipilans, recorded Aug. 25, 1969, at Sempi, Madang Prov., Papua New Guinea; AMNH 83004 (holotype), air 24.00 C., Dept. Herpetology tape no. 190.

en within a relatively short distance of one the fingers." The lectotype and three syn- another. I obtained a pipilans at Wanuma, types of ateles that are in the best condition 700 m., Adelbert Mountains, and a specimen have the finger discs slightly broader than of shellyi (BBM 5788) comes from 15 km. those on the toes, as Boulenger described. north-northwest of Wanuma, 1500-1600 m. In contrast, the toe discs are the broader Comparison with Cophixalus ateles is ones in all specimens of pipilans. The evi- hampered because that species is known dence here too points to the distinctiveness from only the type series of tiny (largest, 15.3 of the two forms. mm. SV), soft, and somewhat faded speci- CALL: Knowledge of the call (fig. 11) is mens. I have examined the lectotype (MSNG based on recordings of three individuals 29116A) and seven syntypes (MSNG 29116B, made at Sempi and 2.5 mi. north-northwest five specimens; BMNH 1947.2.12.6, of Alexishafen, Madang Province, on August 1947.2.12.7). The type locality, "Moroka, 25, September 14, and September 15, 1969. Bartholomew Range, 2300 feet" (Boulenger, The call is a series of rather soft, high- 1898), is south of the Owen Stanley Range, pitched peeps given in groups of 20 to 33 some 300 km. from the closest locality for C. (N = 6 calls) with from 6 seconds to almost pipilans. 2 minutes elapsing between call groups. At Even with allowance for the difficulty of air temperatures of 24.00, 24.70, and 25.60 C., accurately measuring such tiny specimens in the average call rates of the three individuals poor condition, the difference in relative were, respectively, 93.8, 88.6, and 93.2 notes length of hind legs between ateles and pipi- per minute. Approximate lengths of individ- lans (fig. 10) is much greater than might be ual notes in the same order were 0.23, 0.28, expected among populations of one species and 0.24 sec. The dominant frequency was of Cophixalus. The E-N/IN ratio appears to approximately 5300, 4900, and 5300 Hz in differ between the two, also (fig. 10), but the three individuals. The second individual there is overlap in the ranges. Boulenger in the series, with the slowest call rate, long- (1898) described the toes of ateles as having est notes and lowest dominant frequency is "large discs which are smaller than those of the larger (SV 17.1, compared to 16.1 and 408 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 165

16.2 mm.), a possible explanation for the 1610-2160 m. (76009); Tobo, 1450-1650 m. slight difference in its call. No correlations (84509). with temperature are evident over the nar- Previously published records of this row range of temperatures at which record- species are from the Highlands, in the Mt. ings were made. Hagen, Kubor, and Wahgi-Sepik Divide Unfortunately, I do not have a definitive ranges, where specimens came from eleva- tape-recording of Cophixalus shellyi for com- tions of 1520-2900 m. (Zweifel, 1956b, 1962; parison. I have heard (and seen) frogs of this Tyler, 1963a, 1963c). Material reported here species call at two localities in the Western extends the range to the Huon Peninsula and Highlands, and described the call in my field provides intermediate records in the vicinity notes as a "series of short, high-pitched of Wau, Morobe Province (fig. 12): Mt. Mis- peeps . . . [which] may last 10-20 seconds." sim, ca. 10 km. NE. Wau, ca. 1600 m. (BBM What I think is the call of shellyi may faintly 6428, 6433-6436, 6438); Upper Watut River, be heard in the background of a recording of 1100-1600 m. (BBM 6743); Kunai Creek, Edie another species made at one of the Western Creek Road, 1350 m. (AMNH 81148). Highlands localities. An audiospectrogram Frogs of the Huon Peninsula and the High- of this call (air temperature 14.40 C.) shows lands agree in details of coloration, being a note repetition rate of about 340 per min- grayish brown dorsally with the side of the ute, note length of about 0.1 sec., and dom- head black and with peach color prominent inant frequency of 5200 Hz. The great dif- in life in the groin and on anterior and pos- ference in note repetition rate between the terior surfaces of the thighs. They share this call of the new species and the presumed call same color and pattern with Cophixalus pi- of shellyi (it should be even greater at com- pilans. There are some indications of geo- parable temperatures) is evidence for their graphic variation in proportions in C. shellyi specific distinctness. (fig. 10), but the ranges of variation in three ECOLOGICAL NOTES: My specimens samples overlap broadly, in contrast to their came from lowland forest on hillsides at el- relationship to C. pipilans. evations close to sea level, probably no more The specimens I obtained on the Huon than 50 m. One captured in the daytime was Peninsula were collected by children who in leaf litter, whereas those taken at night found them in the daytime (along with Co- were on low shrubs, no more than 1 m. high. phixalus variegatus and Oreophryne sp.) in Few if any frogs called when no rain had the hollows of cut bamboo stems and in the fallen recently, but many were heard on ex- leaf axils of banana plants. cessively humid nights. Tyler (1963a, 1963c) referred specimens of ETYMOLOGY: The specific epithet, this species to C. ateles and considered shel- meaning "peeping," comes from the Latin lyi to be questionably valid. In addition to verb pipilo. the great difference in the E-N/IN ratio (fig. DISTRIBUTION: Cophixalus pipilans is at 10), C. ateles differs in having the toe discs present known only from the vicinity of Lae, slightly broader than those on the fingers, Morobe Province, the coastal region imme- whereas in shellyi only rarely is the disc of diately north of Alexishafen and the Adelbert the third finger as wide as that on the fourth Mountains, Madang Province (fig. 12). The toe; see the foregoing description of C. pi- known range spans about 270 km., but prob- pilans. ably extends both north and south along the coast from the presently known extremes. Cophixalus variegatus (van Kampen) All known localities are given in the foregoing paragraph on paratypes. Tumnang, 1340 m. (74990-74996+4; MCZ 28401); Gang Creek, 1340-1550 m. (74997, Zweifel 76010-76011+2); Mt. Kembiam, ca. 1400 Cophixalus shellyi m., inland of Wasu (BBM 6506, 6507); near Tumnang, 1340 m. (74984-74989); head of Ibang village, 1450 m., inland of Wasu (BBM Kua River Valley northwest of Avengu, 5813, 5814, 5816). 1980 ZWEIFEL: HUON PENINSULA FROGS AND LIZARDS 409

FIG. 12. Distribution of Cophixalus pipilans (open circles), C. shellyi (closed circles), and C. ateles (cross) in Papua New Guinea.

Cophixalus variegatus belongs to a group similar undescribed species exist. Use of the of small scansorial or arboreal species char- name variegatus for the Huon Peninsula acterized by having the fifth toe longer than frogs is tentative and will require review the third. In last reviewing these frogs (Zwei- when the whole group can be studied. The fel, 1962), I recognized only three species type locality of variegatus, Digoel River, Ir- with this characteristic, the others being C. ian Jaya, is about 800 km. west of the Huon darlingtoni Loveridge and C. rostellifer Peninsula. I have collected frogs of the same Wandolleck. The latter has, with good rea- species as those on the Huon Peninsula son, been transferred to a resurrected, (judged by similarity of the calls) on Mt. monotypic genus, Choerophryne (Menzies Kaindi near Wau, Morobe Province, and at and Tyler, 1977). an intermediate locality, Wagau, Herzog Since the publication of my 1962 paper I Mountains. have had the opportunity to tape-record and Frogs that I collected were calling at night study variegatus-group frogs in the field, and from low bushes. Dr. R. G. Hoogland found it is now clear that several morphologically them in the daytime in clumps of fruit grow- 410 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 165

**s, e S

IS- 2 ..rrTrr!rrr!..

T * N..-. --. '. Z A,Si*----3L S. Ak,I...... 55 iL.[LILLLLLL. £ i. .. kjL. C I .2 .4 .6 .8 1.0 1.2 1.4 TIME IN SECONDS FIG. 13. Audiospectrogram (50 Hz filter left, 300 Hz filter right) of call of Oreophryne sp., recorded by S. Grierson on May 27, 1964, at Pindiu, Morobe Prov., Papua New Guinea; AMNH 76030, temperature not recorded, Dept. Herpetology tape no. 204. ing on the trunks of Ficus trees, and other several of the islands southeast of New collectors found them in cut bamboo stems Guinea, on the north coast of Papua New and in banana plants. Guinea and into Irian Jaya remains to be resolved. Copiula fistulans Menzies and Tyler I captured my specimens in the Botanic Lae (81131, 81132, 83032-83036; MCZ Garden at Lae in August 1968 and September A79706); 4 km. N. Lae (103191; MCZ 89221- 1969. All are males which attracted my at- 89227). tention at night with their quiet trilled call Menzies and Tyler (1977) resurrected the (Type 2 call of Menzies and Tyler). Two were in depressions in the soil covered by genus Copiula (formerly synonymized under Cophixalus) and included in it this species, wet leaves, the others in depressions in mowed grass several inches high. None was type locality 11 km. north of Lae. Other lo- truly "subterranean," as characterized by calities include Sattelberg on the Huon Pen- Menzies and Tyler. insula and near Popondetta, Northern Prov- ince. Presumably Mehely's (1901) record of Copiula oxyrhina from Sattelberg represents Hylophorbus rufescens rufescens Macleay this species. Menzies and Tyler dealt only Tumnang, 1340 m. (75018-75022); Gang with Copiula in eastern Papua New Guinea. Creek, Mt. Rawlinson, 1340 m. (75023, Because the three species of Copiula are dis- 75024, 76012-76023); Pindiu, ca. 790 m. tinguished mainly on the basis of their mating (76024); inland (S.) of Wasu Patrol Post, 1450 calls, the systematic status of Copiula on m. (BBM 5812, 5815). 1980 ZWEIFEL: HUON PENINSULA FROGS AND LIZARDS 411

The type locality of a synonym of this growth. Hobart M. Van Deusen and S. 0. common ground frog, Metopostira ocellata Grierson captured individuals at Pindiu that Mehely, is Sattelberg, on the Huon Penin- were calling (fig. 13) from sites 4 to 8 feet up sula (Zweifel, 1972a). in banana and pandanus plants, and I found calling males similarly situated at Lae. I nei- Oreophryne sp. ther found nor heard this species at Gang Creek in the tall primary forest. A photo of Tumnang, 1340 m. (75026-75040+14); Pin- an individual of this species from Lae iden- diu, 820-914 m. (76025-76030); Lae (75041- tified as Oreophryne biroi appears in Zweifel 75044, 81195). (1972d, fig. 5). A user of the key in Parker (1934, pp. 160- 161) could identify these frogs as Oreophryne Phrynomantis lateralis (Boulenger) biroi (Mehely). I hesitate to use that name for these specimens, despite references in Masba Creek, 610 m. (76007, 76008); Lae the literature to biroi from the Huon Penin- (74888, 81058); ca. 24 km. N. Lae, 240 m. sula (Sattelberg: Mehely, 1901, p. 252; Par- (81059). ker, 1934, p. 170, refers to the same speci- Most individuals were calling on the mens as Mehely) because of evidence that ground in rain forest when captured; one was more than one species may currently be cov- under a log. Mehely (1901, p. 220) recorded ered by the one name. this species from Sattelberg. For a photo- The type locality of 0. biroi is "Friedrich- graph, audiospectrograms, and distribution Wilhelmshafen" (= Madang; Mehely, 1897). map see Zweifel (1972a). Near Sempi, about 20 km. north of Madang, I collected specimens of an Oreophryne that Phrynomantis robusta (Boulenger) also keys out to biroi in Parker's key but has Tumnang, 1340 m. (74889). a mating call vastly different from that of the The only specimen of this species captured frogs on the Huon Peninsula and differs as on the Expedition is a juvenile found in a well from them in some proportions not hole in the cut bank of a trail in second treated by Parker. An additional complica- growth forest. Other localities for the species tion is produced by a specimen from Madang within the area of this report are given by itself that, on the basis of its proportions, Zweifel (1972a, p. 499): Aregenang, Mindik, may represent a third species-possibly the Sattelberg, Simbang, and Tewep. true biroi. Unfortunately, I have no information on the mating calls of frogs at Ma- dang. Parker treated several published Sphenophryne mehelyi Parker names as junior synonyms of biroi, but there Mindik, 1200-1600 m. (BBM NG 55310, is no .way at present to justify assignment of field number); 4 km. N. Lae (MCZ 89220). any of these names to populations discussed The type locality of this species is Sattel- here. A frog from Sattelberg identified by berg (Parker, 1934, p. 157). The two speci- Vogt (1911) as Hylella brachypus and by Ty- mens on which Parker based the species ler (1964a) as Oreophryne sp. may represent (specimens referred to Chaparina fusca by the species collected on the Archbold Ex- Mehely, 1901) have been destroyed, and no pedition. others have heretofore been reported from The form found on the Huon Peninsula New Guinea, although Tyler (1967) identified evidently is fairly widely distributed, for I specimens from New Britain as S. mehelyi. have collected it as well as Kokoda (identi- The Huon specimens agree closely with fication based on recordings of mating calls Parker's and Mehely's descriptions. Ventral as well as on morphology). Native collectors color patterns-dusky throat and chest with and I found these frogs at Tumnang hiding small light spots, abdominal area paler with during daytime in banana plants and in cut brown mottling-are virtually identical with bamboo stems in gardens and in second that of one of the typical specimens illus- 412 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 165 trated in color by Mehely (1901, pi. 12, fig. northernmost specimens are from Boana, 40 3). km. west-northwest of Lae (MCZ field nos. I have examined 23 specimens of Sphen- X23502-X23523, X23585). A peculiar aspect ophryne from New Britain, including three of the distribution of S. palmipes is its pres- studied by Tyler and referred by him to S. ence south of the central mountain range, mehelyi. These specimens are all remarkably where Fred Parker obtained numerous spec- similar in having all lower surfaces-chin, imens on the Pio River drainage south of Mt. chest, abdomen, and limbs-uniform brown Karamui (MCZ). The pattern of distribution with small white spots. Thus, they differ described is closely paralleled by that of from the Huon specimens, which agree with another microhylid, Cophixalus cheesmanae Parker's (1934, p. 156) description: "lower (Zweifel, 1979b). surfaces dirty white, the throat and chest Sphenophryne palmipes is riparian, being washed with pale brown and irregularly spot- found beside and in small streams. It is pos- ted with white." The specimen from Lae sibly the only New Guinean microhylid with measures 30 mm. snout to vent, whereas no aquatic habits. The frogs are dull-colored in specimen among 55 in the New Britain series life, greenish brown dorsally with obscure measured by Tyler was as long as 21 mm. In darker markings, and gray beneath with his key to Sphenophryne, Parker (1934, p. darker gray flecks and mottling on the chin. 153) gave "circa 28 mm." as the size of mehelyi, although the type measured 20 mm. (p. Xenobatrachus rostratus (Mehely) 157), and Mehely (1901, p. 257) stated, "das grossere 24.5 mm. lang." As Tyler (1967) Gang Creek, 1340 m. (76048, 76049); di- noted, differences in methods of measuring vide between Ogeramnang and Tobou, 1830- may contribute to the apparent variation. 1980 m. (76050-76052); head of Kua River Differences in size and coloration leave lit- Valley, northwest of Avengu, 1650-2160 m. tle doubt that the specimen from Lae (and, (76053-76098+7); Indagen, north slope of by association, the smaller and less well-pre- Mt. Kiren (76099); near Indum (76100), vic. served specimen from Mindik) belongs to a Ibang Village, south of Wasu, ca. 1450 m. Britain (BBM 6501, 6502). species distinct from the New sam- Two specimens from Gang Creek were ple, which represents an undescribed species. found in leaf litter when trees were being felled for the campsite clearing. Hobart M. Sphenophryne palmipes Zweifel Van Deusen and S. 0. Grierson collected Gang Creek, 1340 m. (75048-75053, several individuals that were calling from 76038-76047); near Zangaren, ca. 1370 m. within the humus during gentle daytime (76031, 76032); Numbut, ca. 1220 m. (76034- rains. 76037); Masba Creek (76033); Tuwap, 1350 Mehely (1901, p. 233) recorded this m. (BBM 1048); Finschhafen (SAM 5684 species from Sattelberg. Zweifel (1972a, pp. [eight specimens]); Ulap, 800-1100 m. (BBM 524-527) considered X. rostratus divisible 5322). into three morphologically and geographical- Originally described on the basis of spec- ly distinct groups, one of which comprised imens from Mt. Dayman and Goodenough the sample from the Huon Peninsula. Island on the southeastern tail of New Guinea (Zweifel, 1956a), Sphenophryne palmipes proves to have an extensive range on Xenobatrachus subcroceus the northern slopes of eastern Papua New Menzies and Tyler Guinea. Tyler and Menzies (1971) reported Ca. 11 km. N. Lae (90767); 4 km. N. Lae it from Alotau, on Milne Bay, the Fifth Arch- (MCZ 89312). bold Expedition obtained specimens on Nor- The first specimen cited is a paratype of manby Island, and I found it at Kokoda, Gar- this recently described species (Menzies and aina and near Wau (Zweifel, 1976). The Tyler, 1977), kindly donated by Mr. Men- 1980 ZWEIFEL: HUON PENINSULA FROGS AND LIZARDS 413 zies. This species is quite distinct from X. ural History, in addition to those cited above rostratus which occurs at higher elevations from Lae and vicinity, establish the presence on the Huon Peninsula, but is exceedingly of the species along the north coast of Papua similar to X. mehelyi (Boulenger), known New Guinea in the Adelbert Mountains and from the southern drainages of Papua New near Alexishafen (Madang Province), at Ma- Guinea. Although Menzies and Tyler de- prik (East Sepik Province), and at Lumi and scribe the call of subcroceus, recordings of Mt. Menawa (West Sepik Province). mehelyi unfortunately are not available for Like the other large Cyrtodactylus, this comparison. Evidence of different vocaliza- species is arboreal and nocturnal, and readily tions would strengthen support for the dis- found by its deep red eye shine in a flashlight tinctness of subcroceus, which now rests beam. Curiously, despite its large size it may largely on geography. be found on saplings scarcely as wide as its body, but it inhabits larger trees also. SAURIA Cyrtodactylus pelagicus (Girard) GEKKONI DAE Lae (66731, 66732, 95177, 95653); 11 km. Cyrtodactylus loriae (Boulenger) N. Lae (103242-103244); Gusiko (66668). Near Pindiu, ca. 910 m. (95647). Earlier records from the Huon Peninsula This dull, gray-brown-colored gecko is include Vogt (1911) and Loveridge (1948). found on tree trunks in forest. The iris is light This adaptable species is widespread over brown, conspicuous against the grayer color the Pacific islands. On New Guinea it is both of the head. terrestrial and arboreal, found on tree trunks at night, and sheltering in cocoa husk piles Cyrtodactylus louisiadensis (de Vis) in the daytime. I have even seen it active on a shaded tree trunk in the daytime, which is Lae and near Lae (66702, 95652); Busu unusual for a Cyrtodactylus. River, 13 km. N. Lae (95169); ca. 24 km. N. Lae (103240); Masba Creek, 610-640 m. (95648-95651). Cyrtodactylus vankampeni (Brongersma) This species, like C. loriae, is typically a tree dweller, although two of the specimens Lae (95654, 103257, 103258). from Masba Creek were found on rocks. So far as I have been able to determine, Others were on tree trunks, including one this species has been reported in the litera- lizard 15 feet above the ground. The eyes ture only once since the original description. reflect a deep red color in the beam of a flash- Brongersma (1933) described it on the basis of two specimens from "the surroundings of light. Modderlust" in former Netherlands New Guinea. De Rooij had earlier (1919) listed the Cyrtodactylus novaeguineae (Schlegel) same specimens as Gymnophthalmus pelag- Lae (95165-95168); Busu River, 13 km. N. icus from "Modderlust am Tami-Fluss" and Lae (95175). "Zwischen Modderlust und Kasawari an der This magnificent gecko (snout-vent length Humboldtbai." Neither Modderlust nor Ka- of the two largest American Museum speci- sawari appears on any map available to me, mens, 172 mm.) is widely distributed in the but the former appears to be no more than lowlands of New Guinea, although rarely about 10 km. south or southeast of Humboldt mentioned in the literature. Brongersma Bay. Loveridge's (1948) record for Aitape (1934) established the validity of the species (Papua New Guinea: West Sepik Province) and gave several localities on the north and is about 180 km. east-southeast of the type south coasts of western New Guinea, as well locality, and the specimens from Lae extend as less specific ones for eastern New Guinea. the range another 680 km. east-southeast. Specimens in the American Museum of Nat- The similarity of C. vankampeni to C. pe- 414 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VO0L. 165 lagicus was noted by Brongersma (1933) and Lepidodactylus lugubris is implicit in de Rooij's confusion of the two (Dumeril and Bibron) species. In the field, the obviously gravid Lae (92316, 92317); Gusiko (66666). condition of such a small lizard (snout-vent This parthenogenetic house-gecko and the length, about 33 mm.) called my attention to similar bisexual species L. novaeguineae oc- the distinctiveness of vankampeni. I found cur sympatrically at Gusiko, where M. Kurtz my two specimens on the ground beneath collected both on trunks of palm trees at broad pieces of banana stems in the Botanic night. Loveridge (1948) reported specimens Garden. from Gusiko and Finschhafen. Gehyra mutilata (Wiegmann) Lepidodactylus magnus Brown and Parker Lae (66707-66711, 92315, 95219, 103261). Lalang, 1400 m. (95655). Loveridge (1948) recorded this species The specimen is a paratype of this recently from Finschhafen and Gusiko. Typically a described species (Brown and Parker, 1977). house-gecko, mutilata sometimes is found in Unlike other Papuan Lepidodactylus, it is a more "natural" habitats, as two of the Lae lizard of interior upland regions, found in the specimens found by M. C. Kurtz in a cave. central mountainous region of Papua New Guinea as well as on the Huon Peninsula. Gehyra vorax Girard Lepidodactylus novaeguineae Lae (95216). Brown and Parker Gehyra vorax was included within the synonymy of G. oceanica by Wermuth (1965, p. Gusiko (66665, 66667). 35), who stated "syn. fide de Rooij 1915." These specimens are paratypes (Brown De Rooij, however, treated oceanica and and Parker, 1977). Loveridge (1948) reported vorax as distinct species. Burt and Burt this species (as L. pulcher) from Gusiko and (1932) considered vorax "probably" a syn- Finschhafen. Records given by Brown and onym of oceanica. I treat vorax as a species Parker outline a distribution at low eleva- on the advice of Mr. and Mrs. Wm. Beckon, tions along the north coast of New Guinea who have accumulated evidence that the two from the area of Jayapura (Hollandia) to the are distinct in sympatry and who examined Huon Peninsula. See remarks under L. lu- the specimen cited. gubris, above. My specimen was on the buttress region of a small tree at night. The species is wide- PYGOPODIDAE spread in the Pacific islands, but there are no Lialis jicari Boulenger other records for the Huon Peninsula. Lae (95659). Gekko vittatus Houttuyn This specimen represents the only record for the Huon Peninsula, although the species Pindiu, ca. 810 m. (95657, 95658); Lae is widespread in lowlands on both north and (66704, 95656). south coasts of New Guinea, and ranges as This widespread gecko of arboreal habits high as 1500 m. (Kluge, 1974). often is found around dwellings, but also occurs in forest at low elevations. AGAMI DAE Hemidactylus frenatus Gonocephalus modestus Meyer Dumeril and Bibron Between Zangaren and Bullum River Lae (92304-92314, 95217, 95218). (95158); Masba Creek, ca. 640 m. (95644). This widely distributed house-gecko does One was found on the ground in second not appear previously to have been recorded growth in the daytime, and another on the from the Huon Peninsula. ground in rain forest at night, but my expe- 1980 ZWEIFEL: HUON PENINSULA FROGS AND LIZARDS 415 rience elsewhere indicates that individuals of This subspecies has been reported only this species are more often in trees. In light from the somewhat indefinite type locality, phase, the dorsal body surface of G. modes- "Sepik-Gebeit" (Mertens, 1931), and from tus is bright green with little or no trace of two localities on the lower Sepik River (He- pattern. An individual can darken, however, diger, 1934). In addition to the specimens so that green is largely replaced by brown, from Lae, I collected a series at Hayfields dark crossbands appear on the body and (10 km. south of Maprik), East Sepik Prov- crossbands faintly evident on the tail in light ince (95248-95254), and the Sixth Archbold phase become much more distinct. The ven- Expedition obtained one (92364) at Oomsis, tral surfaces of the body and limbs are white. 35 km. (by road) west of Lae, Morobe Prov- The iris is brown. ince. In the spelling of the generic name of this These lizards in life have a pale grayish species, I follow Wermuth (1967). brown dorsal ground color with an indistinct paler dorsolateral band on each side, a gold- Gonocephalus papuensis (Macleay) en sheen to the head, and the lining of the mouth black or nearly so. The color pattern Lae (95646, 104861). is thus much as described and illustrated for The genus is in need of revision, therefore pallidus by Mertens (1931), and contrasts the identification is tentative. Vogt (1911) re- with the more distinctly striped condition of corded papuensis from Sattelberg. C. b. novaeguineae of the northwest coast of New Guinea. Loveridge (1948) recorded SCINCIDAE a specimen from Gusiko as novaeguineae, Generic subdivision of the Scincidae, es- but his description suggests an animal with pecially of the subfamily Lygosominae, has indistinct striping. run the gamut from lumping (Smith, 1937) to The lizards taken at Lae were sunning on splitting (Mittleman, 1952). Subsequent con- the trunks of large trees in company with tributions by Greer and by Fuhn have pro- Lamprolepis smaragdina. The color of Cryp- vided reasonable solutions to many of the toblepharus blends well with that of the tree problems of generic assignment. With excep- bark, and the lizards' depressed body form tion made for later work I have followed the enables them to hide readily under loose generic usage of Greer (1970a, 1974). flakes of bark. Carlia fusca (Dumeril and Bibron) Emoia atrocostata irrorata (Macleay) Finschhafen, sea level to 150 m. (95661- Finschhafen, sea level (95676-95682). 95669); Pindiu, 850 m. (95660); Kua River The specimens were collected in the tidal Canyon between Tumnang and Kwenzeng- zone of a coral shore. I find no previous rec- zeng (95221); Gusiko (66679-66681+ 10, ords for this Emoia on the Huon Peninsula, 66688-66690+12). although it is abundant on rocky coral shores Loveridge (1948, pp. 360-365) recognized elsewhere in New Guinea. four races of this widespread species (two on the mainland of New Guinea), whereas For- Emoia baudini verecunda Brown cart (1953, pp. 66-67) could not separate Pindiu, ca. 880 m. (95683); Masba Creek mainland populations on a basis logical for (95684); between Zangaren and Bullum River both geography and morphology. A more (95317, 95319); between Bullum River and thorough study than has been made so far is Selimbeng (95337, 95338). needed before this variable species is prop- These specimens agree closely with E. b. erly understood; hence, I use the binomial. verecunda as described by Brown (1953). The ventral surfaces are relatively pale, the Cryptoblepharus boutonii pallidus sides are dark brown (almost black) with Mertens scattered light spots but without stripes, and Lae (95255-95258, 103292-103297). there is a broad, bronze dorsal band six and 416 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 1 65 two half-scales wide on the body. The scale The captures of caeruleocauda on the counts extend slightly out of the ranges given Huon Peninsula were at moderate to low el- for verecunda by Brown (1953, pp. 4, 6), but evations, as is typical of this species. his sample included only eight specimens. These lizards bear a close resemblance to Emoia cyanogaster longicauda Macleay E. loveridgei, a sympatric species of similar Gusiko (66678). size. The unspotted (but usually striped) Brown (1954, p. 265) identified this speci- sides and heavily barred labial region of lov- men. eridgei serve to distinguish that species from baudini. I noted in living baudini a yellowish tint to the labial region that was not present Emoia loveridgei Brown in loveridgei. Pindiu, 880-910 m. (95312, 95693); Gang Previous records for this species lie on the Creek, 1340 m. (95694, 95707-95718); Kot- north coast of New Guinea. The closest giv- kin, 1220 m. (95695-95698); Zangaren, 1370 en by Brown (1953) to the localities reported m. (95699-95706); between Zangaren and here is Marienberg, about 450 km. north- Bullum River (95314-95316, 95318); between west. Selimbeng and Zangaren (95313); between Bullum River and Selimbeng (95320-95322); Emoia caeruleocauda caeruleocauda between Selimbeng and Tumnang (95339); (de Vis) Tumnang, 1340 m. (95323); between Tum- nang and Kua River (95324, 95325); between Finschhafen, sea level (95688-95692); Pin- Kua diu, 850-880 m. (95685-95687); Gusiko River and Kwenzengzeng (95326, (66687); Lae (66703+3, 66729, 66730, 92327- 95327); near Numbut (95719-95725+6); Gu- 92334, 95274-95276, 103309). siko (66686, 109522, 109523); 2 km. N. Bonga Brown (1956) and Greer (1968) used the (66685). name Emoia callisticta for this species, The majority of these specimens agree which earlier authors (e.g., Loveridge, 1948) closely with the original description (Brown, had called caeruleocauda. I have examined 1953, pp. 10-11). There is a broad, light the holotype of Euprepes callistictus Peters brown dorsal band, the sides are dark brown and Doria, 1878 (MSNG 28049) and find it to and unspotted, and there is a weak light line be, as originally described, a lizard with from axilla to groin, extending anterior to the strongly bicarinate dorsal scales. This is in forelimb in some individuals. Occasional liz- marked contrast to the smooth scutellation ards show scarcely a trace of the lateral light of caeruleocauda, and I do not doubt that line even when alive. the two are different species. The species The specimens from Gusiko and 2 km. that appears closest to Euprepes callistictus north of Bonga are paratypes. They were cit- is Lygosoma acrocarinatum Kopstein, 1926 ed by Brown (1953, p. 10) as AMNH nos. (Emoia acrocarinata: Brown, 1953, p. 23). 66685, 66686 and two uncatalogued. The pre- The two agree in pertinent characteristics of viously uncatalogued specimens have been scutellation, including numbers of scale rows assigned numbers. Brown listed the speci- around midbody, numbers of rows from oc- men from 2 km. north of Bonga as having ciput to rump, number of lamellae under the come from Gusiko, owing to a mistake in a fourth toe, absence of enlarged nuchals, and catalogue entry that has been corrected by the bicarinate nature of the dorsal scales. reference to the collector's field catalogue. The type localities of both species are on the Vogelkop Peninsula of Irian Jaya. I suspect Emoia mivarti (Boulenger) that a more detailed study (I have not seen Pindiu, ca. 850-910 m. (95728-95754+26); the holotype ofacrocarinatum) will establish between Zangaren and Bullum River (95288); that acrocarinatum is a junior synonym of Kabwum, ca. 1370 m. (95755-95757); Lae callistictus. (66726-66728, 103341, 103342). 1980 ZWEIFEL: HUON PENINSULA FROGS AND LIZARDS 417

Emoia mivarti is somewhat of a reptilian (95771); Numbut (95776, 95777); Zangaren, weed, often abundant in disturbed habitats 1370 m. (95778); between Zangaren and Bul- but scarcely, if at all, penetrating mature for- lum River (95343, 95344); between Bullum est. The absence of specimens from the Gang River and Selimbeng (95328, 95336); Tum- Creek camp in contrast to its abundance at nang, 1340 m. (95330); Kua River Canyon, Pindiu reflects this habitat preference. I have between Tumnang and Kwenzengzeng not found previous references to this species (95329); Lae (66724, 66725). on the Huon Peninsula, but probably speci- These are brown skinks with little contrast mens were referred to other species. in the color pattern. One adult female had, in life, a darker brown dorsolateral streak Emoia pallidiceps pallidiceps de Vis bordered below by lighter brown, the same Between Selimbeng and Tumnang (95340, as that of the back. Another adult female had 95341); Gang Creek, 1370 m. (95342, 95763- the sides slightly darker than the back and 95767); Zangaren, 1370 m. (95758-95762); spotted with grayish white, but without clear Kotkin, 1220 m. (95768, 95769). demarcation between the color of the sides These specimens have dark, unspotted and that of the back. Lateral spotting is more sides and a light lateral line that passes from distinct in juveniles, especially on the neck. the ear to the groin. This is the pattern char- In both juveniles and adults the head has a acteristic of pallidiceps as Brown (1953) de- slight bronze tint, the labial scales are dull fined this species. The mean number of yellow and the ventral surfaces gray. An scales around midbody in eight specimens adult had yellow eyelids. from the Huon Peninsula is 33.1 (range 32 to Brown (1953) recorded the specimens 34), and so by Brown's criteria the speci- from Lae. I assume that Loveridge's (1948) mens represent the nominate subspecies record for E. tropidolepis from Gusiko refers rather than E. p. maxima, which has a range to this species, since Brown considered trop- in this scale count of 38 to 42. Several spec- idolepis a subspecies ofphysicae. imens from the Huon Peninsula exceed the maximum snout to vent length of 52 mm. giv- Emoia submetallica popei Brown en by Brown forpallidiceps; the largest mea- Masba Creek (95781); Lae (66705). sures 59 mm. The color pattern of the lizard from Masba A specimen from Gang Creek (95342) had Creek agrees with Brown's (1953) descrip- the following colors and pattern in life: top tion of Emoia s. popei, except that a trace of of head bronze, fading along back to greenish a light line is present between the ear and the brown in lumbar region; a narrow dorsolat- groin. There is no light line on the side of the eral light line faintly indicated, strongest on body, only obscure spotting on the dark the neck; a black lateral band bordered be- field. The number of scale rows around mid- low by a more distinct light line running from body (36) and number of fourth toe lamellae ear to groin with a branch onto anterior side (40) are relatively high, as is characteristic of of forearm; ventral surfaces, including un- this form, and help to distinguish it from the derside of tail, rust-colored; iris bronze. somewhat similar species E. baudini and E. Brown (1953) cited no records for this loveridgei which occur in the same general species on the Huon Peninsula, but he ex- region. amined specimens from areas to the north- Brown (1953) recorded E. s. popei from west, west and south. localities to the northwest and southeast of the Huon Peninsula, but not from the Pen- Emoia physicae physicae insula proper. (Dumeril and Bibron) Masba Creek, 610 m. (95772-95775); Kot- Eugongylus rufescens (Shaw) kin, 1220 m. (95770); Gang Creek, 1340 m. Lae (103490, 103491). 418 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 165

Loveridge (1948) recorded this large, This species was known under the substi- ground-dwelling skink from Finschhafen. tute name elegantoides Ahl 1925 for many years; Greer (1974) restored elegans. When Lamprolepis smaragdina perviridis Loveridge (1945) described Lygosoma ele- (Barbour) gantoides lobulus, he had available only the type series of 16 specimens from Mt. Wil- Finschhafen, sea level (95670-95673+8); helm. He knew the typical subspecies only Gusiko (66684); Lae (66722, 66723, 92320, from Boulenger's (1897) original description 92321, 94559, 95259, 95260, 95674). of the holotype from Mt. Victoria and from The Archbold Expedition found this com- additional information on that specimen and mon tree skink only at the lowest-elevation one from Moroka communicated by Mal- camp. It is common in the Botanic Garden colm Smith (Loveridge, 1945, pp. 49-51). at Lae, where clearing has exposed the The new form was distinguished by having trunks of large, forest-grown trees to sun- more scale rows around midbody (34 to 36 light. I follow Greer (1970b) in the use of the as opposed to 32) and by having the distal generic name Lamprolepis, rather than the subdigital lamellae "not or scarcely differ- more familiar Dasia, which he restricts to entiated from those of the basal" part of the other species. toe (more marked differentiation in the typical form). A specimen in the American Mu- Lipinia longiceps (Boulenger) seum of Natural History (59046) is from Mt. Gusiko (66697); Lae (92326). Tafa, 2070 m., about 80 km. west-northwest Melvin C. Kurtz found the Gusiko lizard of the type locality of elegans. It has 36 on a tree trunk, which agrees with Stickel's scales around midbody and 19 subdigital la- observations (Loveridge, 1948, p. 357). For mellae that show essentially the same a photograph, distribution map, and key amount of differentiation as in topotypic lob- characters of this species see Zweifel ulus with which I compared it. I do not re- (1979a). gard lobulus as sufficiently well characterized, so I use the binomial for the Huon Lipinia noctua (Lesson) lizards. Lobulia elegans, although mentioned in- Finschhafen (95802). frequently in the literature, is a common I have elsewhere (Zweifel, 1979a) reported species at moderate to high elevations all on the variation and geographic distribution along the mountainous backbone of New of this species and have concluded that Ly- Guinea. The specimens collected on the gosoma miotis Boulenger is a junior syn- Archbold Expedition evidently are the first onym of L. noctua. Other locality records found on the Huon Peninsula. Those from for the Huon Peninsula include Lae and Gu- Mt. Ulur were in grassland or at the grass- siko (Zweifel, 1979a). land-forest boundary; one was in a hollow Lipinia pulchra (Boulenger) tree-fern log. Lae (66703, 131175-131179). Lobulia stanleyana (Boulenger) This appears to be the only record for the Gang Creek, 1340 m. (95803-95817); ridge species on the Huon Peninsula, although the north of Gang Creek, 1580 m. (95818); Kot- species is widespread at low elevations on kin, 1220 m. (95819-95821); Kabwum, ca. the north coast of New Guinea. See Zweifel 1370 m. (95822-95825); between Selimbeng (1979a) for a distribution map, photograph, and Tumnang (95413, 95414, 95416); between and discussion. Selimbeng and Zangaren (95415); Gevak, 1460 m. (93009-9301 1). Lobulia elegans (Boulenger) Loveridge (1948, pp. 358-359) considered Gang Creek, 1340-1370 m. (95784, 95785); Leiolopisma morokana Parker (1936) a sub- Mt. Ulur, 2380-2440 m. (95786-95800). species of L. stanleyana. I have elsewhere 1980 ZWEIFEL: HUON PENINSULA FROGS AND LIZARDS 419

(Zweifel, 1972b) given reasons for treating L. The six specimens from the Huon Penin- morokana as a separate species, so I use the sula are the first to be recorded from there. binomial here. The number of dorsal scales ranges from 67 Lobulia stanleyana is a common species to 76 (mean, 71) and so does not agree with in primary forest throughout the entire cen- paniaiense orflavipes as diagnosed by Bron- tral mountainous region of New Guinea. It gersma. The palpebral disc is small but un- has not previously been reported from the divided. Whether any geographic popula- Huon Peninsula, undoubtedly because of the tions of this species deserve nomenclatural dearth of herpetological collections from recognition can be determined only by study higher elevations there. The specimens from of the large amount of material of this species Kotkin (1200 m.) are from the lowest eleva- that has accumulated in museums since tion I know of for this species. Principally a Brongersma's work was done. For the pres- forest-dweller, L. stanleyana occurs above ent, I prefer not to give a subspecific assign- the forest growth, as on Mt. Hagen (Western ment to the specimens from the Huon Pen- Highlands Province) where E. Thomas Gil- insula. liard collected specimens (71200-71225) "on This species is notable for great individual steep grassy slope leading to summit [3440 variation in color and pattern, evidently a m.]" (field notes). polymorphic condition (Greer and Raizes, Lobulia stanleyana and L. elegans fre- 1969; Woodruff, 1972). Two specimens in the quently are found in sympatry. The former present series have a broad brown dorsal typically is seen in sunny spots on the leaf band, eight scale rows wide, that contrasts litter, whereas the latter is more often off the with the light tan of the sides. The remaining forest floor on rocks, stumps, or low on tree four lizards have a pattern on the body of trunks. dark brown crossbands on a slightly lighter ground color. One of the crossbanded spec- Prasinohaema flavipes (Parker) imens had, in life, a gray-green ground color Kabwum, ca. 1370 m. (95826-95829); on the body with dark brown crossbands. Gang Creek, 1370 m. (95831); Kotkin The top of the head was bronze, the iris (95830). brown, the palms and soles bright yellow, Brongersma (1949) described "Lygosoma and the tongue and lining of the mouth blue. (Leiolepisma) flavipes paniaiense" from the The color of the mouth and tongue results Wissel Lakes region of Irian Jaya, and di- from the blue-green color of the lizard's agnosed it as differing from L. flavipes Par- blood (Greer and Raizes, 1969). ker "in having a distinct (although some- This species has suffered more than many times divided) palpebral disc, a slightly others from instability in its generic place- higher number of supraciliaries, and in hav- ment. Originally described as a Lygosoma by ing a higher number of scales from the pari- Parker (1936), it was placed in Leiolopisma etals to the posterior border of the hind- by Loveridge (1948) and in Sphenomorphus limbs." The dorsal scale counts given are 70 by Mittleman (1952). Greer (1970a) listed it to 88 for 23 specimens of paniaiense and 58 as a member of theflavipes species group of to 66 for three specimens of flavipes from Leiolopisma, but earlier, Greer and Raizes Mt. Wilhelm. Parker (1936, p. 89) described (1969) indicated that generic status was war- the holotype as having "no palpebral discs." ranted for flavipes and two related forms. Loveridge (1948, p. 354) described the "dis- Adding to the confusion, Maderson (1970) tinct, though small" palpebral discs of the used Greer's unpublished manuscript name specimens from Mt. Wilhelm, and Brongers- Chlorohaemus for L. flavipes and L. virens. ma referred to a specimen from this locality Finally, Greer (1974) provided the name used as "showing a small disc, although it is more here. granular than in paniaiense." Brongersma suggested that the specimens from Mt. Wil- Prasinohaema virens virens (Peters) helm may represent a third subspecies. Gusiko (66693-66696, 66698). 420 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 165

Loveridge (1948) reported specimens from side), two median rows of dorsal scales Gusiko. slightly broadened, 95 to 109 (mean 101.5; N = 10) scale rows from occiput to base of tail, Sphenomorphus anotus Greer 34 to 36 (mean 34.9; N = 10) scale rows around midbody, the sixth (nine individuals) Masba Creek (95880, 95881). or the fifth (three individuals) supralabial di- At the time of its description (based on rectly beneath the eye, and 20 to 25 (mean specimens collected on the Seventh Arch- 22.6; N = 10) keeled lamellae beneath the bold Expedition; Greer, 1973), this species fourth toe, which is notably longer than the was known only from the type locality. How- third. ever, Dr. Greer informs me (in litt.) that sub- The dorsal ground color is gray-brown to sequently the Museum of Comparative Zo- dark brown with small light gray or light ology has received specimens collected at brown spots arranged roughly in transverse Popondetta, Lego, and Sangara, all in North- rows. The abdomen was yellowish green in ern Province. a juvenile, but peach color in adults. The un- What appears to be the closest relative of derside of the tail is heavily pigmented with Sphenomorphus anotus is S. microtympanus dark gray, the chin and throat somewhat less Greer (1973). It was found at Garaina, about so. The three largest specimens measure 81, 200 km. south of Masba Creek and 160 km. 84 and 85 mm. snout to vent. northwest of Popondetta. These specimens are the first of their species to be identified from the Huon Pen- Sphenomorphus derooyae (Jong) insula. Masba Creek (95867); Gang Creek (95504, 95868-95874); Kabwum (95875-95877). Sphenomorphus granulatus (Boulenger) These are secretive skinks of the forest Masba Creek (95782, 95783). floor. I found one beneath a log, and others The specimens from the Huon Peninsula were captured when they wandered out of were taken on the ground in rain forest. They the leaf litter and onto adjacent bare ground agree in all pertinent characteristics with in the camp clearing. Boulenger's (1903) description and illustra- Identification of the specimens as this tion of the type-specimen, supposedly cap- species, which has scarcely been mentioned tured at an elevation of 6000 feet in the Al- in the literature since its original description bert Edward Mountains, but possibly from (deJong, 1927), is based on comparison with a lower elevation (see Zweifel, 1979a, p. 16). specimens identified as derooyae by Greer I find only three literature references to and on the concordance between characters specimens of this species since the original of scutellation of my sample and of a sample description. Vogt (1912) mentioned, without reported by Greer and Parker (1974, table 2). comment, eight specimens from an indefinite Eight of the 12 specimens in my sample are locality in the interior of Irian Jaya. In a later larger than the maximum of 71 mm. snout- paper (1932) he recorded eight additional vent length given by Greer and Parker (my specimens from the Sepik region, but again maximum, 85 mm.), but this could merely without specific locality. Greer (1970a, p. reflect geographic variation or the larger 172) examined the skull of one of the Masba sample size. Creek specimens. This is an elongate, relatively short-legged form with the following scutellation: no su- pranasals, prefrontals in contact, four supra- Sphenomorphus jobiensis (Meyer) oculars, paired frontoparietals, parietals en- Finschhafen, sea level (95832); Masba closing a large interparietal, eyelid without Creek, 610 m. (95836-95845); Pindiu, ca. a disc, two to five enlarged nuchals on a side 850-910 m. (95846-95858+5); between Pin- (only one specimen has two, and only on one diu and Kwenzenzeng, 940 m. (95474); Zan- 1980 ZWEIFEL: HUON PENINSULA FROGS AND LIZARDS 421 garen, 1370 m. (95833, 95834); Numbut, 1220 Sphenomorphus solomonis (Boulenger) m. (95835); Gusiko (66692, 95923); Lae Lae (66682, 66683+1, 66712-66721, 92325, (66700, 66701+ 1, 92322, 95475, 103609- 103616, 103617); Pindiu, ca. 910 m. (95862- 103611). 95865); Zangaren (95866); Tumnang (95493, This abundant and widespread species has 95495-95500); between Zangaren and Bul- suffered much nomenclatural confusion. lum River (95454). Loveridge (1948, p. 346) recorded specimens The dorsal color, recorded for specimens from the Huon Peninsula as Lygosoma me- from the last two localities listed above, is gaspila papuense and used the name Lygo- olive-brown with little trace of pattern ex- soma variegatum jobiense for the form now cept for slight dark edging to the scales. The known as Sphenomorphus stickeli melano- head tends pleurus. Inger (1958) described S. melano- to be reddish brown, most mark- pleurus from the north coastal region of New edly so in the snout region ofjuveniles. The Guinea chin is gray, the chest generally yellow without reference to Loveridge's pa- changing gradually to peach color in the re- per, but later (1961) he corrected this over- gion of the hind legs. Two juveniles were en- sight and recognized melanopleurus as a sub- tirely gray beneath. species of Loveridge's Lygosoma variegatum This is one of the more abundant ground- stickeli, which he raised to specific rank as dwelling species of low elevations; Greer and Sphenomorphus stickeli. He diagnosed the Parker (1974) map its distribution. specific differences of Sphenomorphus stick- Loveridge (1948, pp. 350, 353) recorded eli and Sphenomorphus jobiensis but failed both "Lygosoma (Sphenomorphus) pardale to note that jobiensis was the same species moszkowskii Vogt" and "Lygosoma (Lygo- Loveridge had identified as L. m. papuense. soma) solomonis schodei Vogt" from the Another contribution to the confused syn- Huon onymy of S. jobiensis was Loveridge's Peninsula. The scale counts and notes (1948, p. 348) treatment of "Lvgosoma on color in life he gave (recorded by W. H. (Sphenomorphus) aruense (Doria)," the Stickel) show no obvious differences be- name he applied to lizards of the Highlands tween the forms. I examined three of the four of New Guinea since named Sphenomorphus specimens Loveridge recorded as moszkow- leptofasciatus by Greer and Parker (1974). I skii, and regard two from Finschhafen and have examined the four syntypes ofEumeces Sorong to be the same species as the lizards aruensis Doria (Museo Civico di Storia Nat- I here identify as S. solomonis. Greer and urale di Genova no. 27901) and am con- Parker (1974) do not consider schodei a valid vinced that Boulenger (1887, p. 247) was cor- subspecies. rect in considering aruensis a junior synonym of Sphenomorphus jobiensis. Sphenomorphus stickeli stickeli Sphenomorphus jobiensis is found in low- (Loveridge) land rain forest and evidently only sparingly as high as recorded here (4000-4500 ft.). Its Masba Creek (95861); Gusiko (109524- absence from collections made at Gang 109529); ca. 2 km. N. Bonga (66691, 95922). Creek, a cool and heavily forested location For a discussion of some of the nomencla- at an elevation of 4500 feet, may indicate that tural confusion surrounding this species, see it occurs at higher elevations only where the the foregoing account of Sphenomorphus jo- mature forest has been opened somewhat. biensis. The specimen from Masba Creek was shot on a log 4 feet above the ground in Sphenomorphus minutus (Meyer) rain forest. One of the specimens from Gu- siko (which are topotypes) was found "on Finschhafen (66995); Masba Creek (95859, coral at bank of stream in jungle. Often seen 95860); Tumnang, 1340 m. (95501). A speci- climbing on coral to bask in the sun" (field men from Masba Creek was found on the notes of M. C. Kurtz; "coral" presumably ground in rain forest. refers to coral limestone outcrops). 422 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 165

Tiliqua scincoides gigas (Schneider) are the easternmost in the distribution of Tri- Lae (95879). bolonotus along the north coast of New Loveridge (1948) recorded this widespread Guinea. species from Finschhafen and Gusiko, and Vogt (1911) had specimens from Sattelberg VARANIDAE and Lialun. Varanus indicus indicus (Daudin) Near Pindiu, 910 m. (95882). Tribolonotus gracilis de Rooij Loveridge (1948) recorded this species Lae (92318, 95438, 103625-103627). from Gusiko, and Vogt (1911) from Bukaua. Vogt (1911) recorded T. novaeguineae It ranges widely over New Guinea. Schlegel from Bukaua on the Huon Penin- sula. The status of gracilis as a species dis- Varanus prasinus prasinus (Schlegel) tinct from novaeguineae is questionable Vicinity of Lae (92335-92337). (Zweifel, 1966), and it is doubtful whether This beautiful arboreal monitor is wide- two species of Tribolonotus occur on the spread in New Guinea. There is an earlier Huon Peninsula or anywhere on mainland record for Sattelberg on the Huon Peninsula New Guinea. The Huon Peninsula localities (Mertens, 1942).

BIOGEOGRAPHY THE GEOLOGIC SETTING area in Middle Miocene. Jaques and Robin- The Finisterre and Saruwaged ranges-the son (1977, p. 295) stated "the northern Fin- mountainous backbone of the Huon Penin- isterre Range-Huon Peninsula region ... sula-are among the results of the long-con- was not uplifted until the Pliocene. Uplift tinued interaction between the Australian since then has been rapid: some 3000 m of tectonic plate on the southwest and the Pa- uplift has occurred since the late Pliocene." cific plate on the northeast that produced the The region is still tectonically active. Studies main island of New Guinea (Dow, 1977). of terraces on the north side of the Huon These ranges, together with other lesser Peninsula indicate rates of uplift ranging ranges of mountains along the north coast of from 0.5 m. to 3.0 m. per 1000 years during New Guinea, are landlocked parts of a Pa- the last 220,000 years (Chappell, 1974). For leogene island arc system of which New Brit- part of the time while the ranges were being ain, New Ireland, and other associated is- uplifted, they remained isolated from the lands still form free-standing segments mainland. In his discussion of Pliocene to (Jaques and Robinson, 1977). From the bio- Holocene sedimentation, Dow (1977) said geographic standpoint, the most significant that "the Ramu-Markham Fault Zone was a aspects of the Huon Peninsula region are its narrow arm of the sea in which was depos- relatively late development in geologic time, ited up to 2000 m of sandstone and con- its isolation from the mainland of New glomerate...." Just when a dryland con- Guinea, and the effects of climatic change nection between the Peninsula and the rest during the Pleistocene. of New Guinea formed does not appear to The collision between plates evidently have been established; it may have been as proceeded from west to east, with the Huon late as the period of lowered sea level in the Peninsula region being uplifted relatively first glaciation. late. Dow (1977, pl. 7) showed no land in the The higher reaches of the Finisterre and 1980 ZWEIFEL: HUON PENINSULA FROGS AND LIZARDS 423

Saruwaged ranges were glaciated in the scarcely more satisfactory, with only two Pleistocene. Loeffler (1971) illustrated glacial specimens now considered to represent the features of the landscape and placed the species (see foregoing species account). Pleistocene snowline at 3650 to 3700 m. He If evidence for endemism is weak among noted that this is some 100 m. higher than on frogs, it is nonexistent among lizards. Suffi- some mountains in the central ranges at the ciently close attention might reveal differ- same latitude, and attributed the difference ences on a subspecific level, but all forms to post-Pleistocene uplift of the Peninsula now recorded are readily referred to species rather than a different local climate. In a later also found off the Peninsula. paper Loeffler (1972, p. 33) stated: "The Pleistocene climate in the mountainous area DISTRIBUTIONAL RELATIONSHIPS was relatively uniform and temperatures WITHIN NEW GUINEA were about 5 to 6° C lower than at present.... Excluding the introduced Bufo and the Pleistocene climate in Papua and New three possible endemics, there remain 34 Guinea was little different from the present- species of frogs to be considered. Nineteen day climate except for the temperatures." At of these are widely distributed at low eleva- the peak of the most recent glaciation (the tions. Distributions of most of the remaining only one for which there is firm evidence), 15 species probably are disjunct across the montane grassland was greatly expanded, low but not to the extent that cool, montane hab- Ramu-Markham Valley. itats Huon Peninsula Evidence for disjunctions of ranges is, of connected the ranges course, basically negative-the absence of with the Bismarck Range across the Ramu- collection records in the intervening regions. Markham Valley (Hope and Hope, 1976). In many instances, however, information on Deglaciation took place some 11,000 to habitat preferences or apparent altitudinal 15,000 years ago (Hope and Peterson, 1975). limitations (possibly reflecting temperature requirements) strengthens the case. Several FAUNAL DISTRIBUTIONAL of the apparently disjunct frog species are RELATIONSHIPS known or assumed to breed in mountain streams of a turbulent, even torrential na- ENDEMISM ture, and the Ramu-Markham Valley pro- Table 1 lists 38 species of frogs (one intro- vides no suitable breeding habitat. These duced) and 52 species of lizards recorded include four species of Nyctimystes and from the Huon Peninsula. A curious aspect Litoria angiana, L. micromembrana, and L. of the fauna is the scarcity of endemic wollastoni, all hylids. The distribution of species, considering the range of habitats Lechriodus aganoposis is poorly known, but available and the isolation of the mountain- no specimen has come from below 1200 m. ous areas of the Huon Peninsula. Among the Three species of microhylids, Cophixalus frogs, only three of 37 native species appear biroi, C. shellyi, and C. variegatus appear to to be endemic, and these are poorly known. be similarly limited. Two species of micro- Nyctimystes obsoleta, known from a single hylids, Sphenophryne palmipes and Xeno- specimen from Simbang, a coastal locality rhina doriae, may be disjunct across the near Finschhafen, appears to be distinct Ramu-Markham Valley, but occur at low el- from any other known Nyctimystes (Tyler, evations in hilly areas elsewhere. Xenobatra- 1965), but confirmation through the study of chus rostratus cannot be classified at pres- additional specimens is greatly needed. ent: the peninsular population appears to be Sphenophryne polysticta is now known only distinctive and restricted to high elevations, from the original description (Parker, 1934) yet elsewhere (as nearby as the type locality, and from Mehely's (1901) illustrations, the Astrolabe Bay) frogs referred to this species type-specimen having been destroyed. evidently occur at sea level. The Huon Rana Knowledge of Sphenophryne mehelyi is grisea probably represent a disjunct popu- 424 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 165

TABLE 1 Frogs and Lizards Recorded from the Huon Peninsula

Family and SpecieSs Source or Authoritya Bufonidae Bufo marinus (Linnaeus) AMNH Leptodactylidae Lechriodus aganoposis Zweifel Archbold Expedition (AMNH) Hylidae Litoria amboinensis (Horst) AMNH Litoria angiana (Boulenger)b Archbold Expedition (AMNH) Litoria eucnemis (Lonnberg) AMNH Litoria genimaculata (Horst) AMNH Litoria i. infrafrenata (Gunther) Archbold Expedition (AMNH) Litoria micromembrana (Tyler)r Archbold Expedition (AMNH) Litoria nigropunctata (Meyer) Menzies, 1972 Litoria thesaurensis (Peters) Archbold Expedition (AMNH) Litoria wollastoni (Boulenger) Archbold Expedition (AMNH) Nyctimystes disrupta Tylee Archbold Expedition (AMNH) Nyctimystes foricula Tylee Archbold Expedition (AMNH) Nyctimystes kubori Zweifelb Archbold Expedition (AMNH) Nyctimystes obsoleta (Lonnberg) Tyler, 1965 Nyctimystes pulchra (Wandolleck)b Archbold Expedition (AMNH) Nyctimystes Sp.b Archbold Expedition (AMNH) Ranidae Platymantis papuensis Meyer Archbold Expedition (AMNH) Rana daemelii (Steindachner) Archbold Expedition (AMNH) Rana grisea van Kampenb Archbold Expedition (AMNH) Rana sp. (pond form) Archbold Expedition (AMNH) Rana sp. (stream form) Menzies, 1976 Microhylidae Cophixalus biroi (Mehely) Archbold Expedition (AMNH) Cophixalus cheesmanae Parker AMNH, SAM Cophixalus pipilans, new speciesb AMNH, MCZ, YPM Cophixalus shellyi Zweifelb Archbold Expedition (AMNH) Cophixalus variegatus (van Kampen) Archbold Expedition (AMNH) Copiula fistulans Menzies and Tyler AMNH Hylophorbus r. rufescens Macleay Archbold Expedition (AMNH) Oreophryne sp. Archbold Expedition (AMNH) Phrynomantis lateralis (Boulenger) Archbold Expedition (AMNH) Phrynomantis robusta (Boulenger) Archbold Expedition (AMNH) Sphenophryne mehelyi Parker BBM, MCZ Sphenophryne palmipes Zweifelb Archbold Expedition (AMNH) Sphenophryne polysticta (Mhely) Parker, 1934 Xenobatrachus rostratus (Mehely) Archbold Expedition (AMNH) Xenobatrachus subcroceus Menzies and Tyler AMNH, MCZ Xenorhina doriae (Boulenger) Zweifel, 1972a Gekkonidae Cyrtodactylus loriae (Boulenger) Archbold Expedition (AMNH) Cyrtodactylus louisiadensis (de Vis) Archbold Expedition (AMNH) 1980 ZWEIFEL: HUON PENINSULA FROGS AND LIZARDS 425

TABLE 1-( Continued) Family and Species Source or Authority'

Cyrtod(actylus novaeguinea (Schlegel) Archbold Expedition (AMNH) Cyrtodactylus pelagicus (Girard) AMNH Cyrtodactylus vankampeni (Brongersma) Archbold Expedition (AMNH) Gehyra interstitialis Oudemans Vogt, 1911 Gehyra mutilata (Wiegmann) Archbold Expedition (AMNH) Gehyra vorax Girard AMNH Gekko vittatus Houttuyn Archbold Expedition (AMNH) Hemidactylus frenatus Dumeril and Bibron Archbold Expedition (AMNH) Hemiphyllodactylus t. typus Bleeker Loveridge, 1948 Lepidodactylus lugubris (Dumeril and Bibron) Archbold Expedition (AMNH) Lepidodactylus magnus Brown and Parker Archbold Expedition (AMNH) Lepidodactylus novaeguineae Brown and Parker AMNH Pygopodidae Lialis jicari Boulenger Archbold Expedition (AMNH) Agamidae Gonocephalus dilophus (Dumeril and Bibron) Lonnberg, 1900 Gonocephalus modestus Meyer Archbold Expedition (AMNH) Gonocephalus papuensis (Macleay) Archbold Expedition (AMNH) Scincidae Carlia fusca (Dumeril and Bibron) Archbold Expedition (AMNH) Cryptoblepharus boutoni pallidus Mertens Archbold Expedition (AMNH) Emoia atrocostata irrorata (Macleay)p Archbold Expedition (AMNH) Emoia baudinii verecunda Brownb Archbold Expedition (AMNH) Emoia c. caeruleocauda (de Vis) Archbold Expedition (AMNH) Emoia cyanogaster longicauda Macleay AMNH Emoia loveridgei Brown Archbold Expedition (AMNH) Emoia mivarti (Boulenger)b Archbold Expedition (AMNH) Emoia p. pallidiceps de Visb Archbold Expedition (AMNH) Emoia physicae physicae (Dumeril and Bibron) Archbold Expedition (AMNH) Emoia submetallica popei Brownb Archbold Expedition (AMNH) Eugongylus albofasciolatus (Gunther) Vogt, 1911 Eugongylus rufescens (Shaw) Archbold Expedition (AMNH) Lamprolepis smaragdina perviridis (Barbour) Archbold Expedition (AMNH) Lipinia longiceps (Boulenger) Archbold Expedition (AMNH) Lipinia noctua (Lesson) Archbold Expedition (AMNH) Lipinia pulchra (Boulenger) Archbold Expedition (AMNH) Lobulia elegans (Boulenger)b Archbold Expedition (AMNH) Lobulia stanleyana (Boulenger)b Archbold Expedition (AMNH) Prasinohaema flavipes (Parker)b Archbold Expedition (AMNH) Prasinohaema v. virens (Peters) Loveridge, 1948 Sphenomorphus anotus Greer Archbold Expedition (AMNH) Sphenomorphus derooyae (Jong)b Archbold Expedition (AMNH) Sphenomorphus granulatus (Boulenger)b Archbold Expedition (AMNH) Sphenomorphus jobiensis (Meyer) Archbold Expedition (AMNH) Sphenomorphus minutus (Meyer/i Archbold Expedition (AMNH) Sphenomorphus muelleri (Schlegel) Vogt, 1911 Sphenomorphus pratti neuhaussi (Vogt) Vogt, 1911 426 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. I165

TABLE 1-( Continued) Family and Species Source or Authoritya Sphenomorphus solomonis (Boulenger) Archbold Expedition (AMNH) Sphenomorphus stickeli stickeli (Loveridge) Archbold Expedition (AMNH) Tiliqua scincoides gigas (Schneider) Archbold Expedition (AMNH) Tribolonotus gracilis de Rooij Archbold Expedition (AMNH) Varanidae Varanus i. indicus (Daudin) Archbold Expedition (AMNH) Varanus p. prasinus (Schlegel) Archbold Expedition (AMNH) a Where specimens were collected on the Archbold Expedition, this is indicated. References for specimens examined including those collected on the Expedition are: AMNH (American Museum of Natural History); BBM (Bernice P. Bishop Museum); SAM (South Australian Museum); MCZ (Museum of Comparative Zoology, Harvard University); YPM (Yale Peabody Museum). Where no specimens were examined, the most pertinent literature reference is cited. b First published record for the Huon Peninsula; records for some other species collected for the first time on the Huon Peninsula by the Archbold Expedition have been published elsewhere.

lation, though there is possible confusion south and west (Zweifel, 1979b). Spheno- with a lowland form (see foregoing account). phryne palmipes is an oddity (fig. 14K). It is The lizard fauna of the Huon Peninsula is found along the north flanks of the eastern composed almost wholly of species wide- ranges, occurs close to sea level at Milne spread at low elevations (though many range Bay (and possibly also at Finschhafen, if that up into the mountains). Only four of the 52 locality is to be taken literally), and has a species are upland forms almost certainly disjunct population in the southern drainages disjunct across the Ramu-Markham Valley: west of its major distributional area. Xeno- Lepidodactylus magnus (Gekkonidae); rhina doriae (not mapped) is distributed Lobulia elegans, L. stanleyana, and Prasi- much like S. palmipes, including a possibly nohaema flavipes (Scincidae). A fourth disjunct western population, but is found on species of skink, Sphenomorphus brunneus, both northern and southern slopes of the is recorded from Boana to the west of the southeastern mountains. Cophixalus varie- Huon Peninsula and probably has its range gatus and Rana grisea (not mapped) are not interrupted by the Valley (Greer and Parker, amenable to analysis because of unresolved 1974). problems with sibling species. Figure 14 illustrates the distributions of Although it is possible to identify an up- most of the species of frogs and lizards with land assemblage of species in the Huon Pen- disjunct populations on the Huon Peninsula. insula disjunct from populations elsewhere, One relationship is immediately evident: al- this is not a fauna restricted to high eleva- most all the species occur in the mountain tions. Lower elevational limits for most of mass west of the Peninsula (see fig. 12 for the species are in the vicinity of 1100-1200 elevations), and some of these extend into m., with none higher than 1400 m. At least the mountains along the southeastern tail of five species (Litoria wollastoni, Nyctimystes New Guinea. pulchra, Cophixalus biroi, Sphenophryne Only one clearly north coast distribution palmipes, Lobulia elegans) are known below is seen-that of Cophixalus biroi (fig. 141). 1000 m., although this may not be character- But even in this instance, the most closely istic. In fact, one of the remarkable features similar species are in the mountains to the of the frog fauna of the peninsular mountains FIG. 14. Distributions in Papua New Guinea of frogs and lizards with disjunct populations on the Huon Peninsula. A. Lechriodus aganoposis; B. Litoria angiana; C. L. micromembrana; D. L. wollastoni; E. Nyctimystes disrupta; F. N. foricula; G. N. kubori; H. N. pulchra; I. Cophixalus biroi; J. C. shellyi; K. Sphenophryne palmipes; L. Lepidodactylus magnus; M. Lobulia stanleyana; N. Prasino- haemaflavipes; 0. Lobulia elegans. 428 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 1 65 is the apparent absence of high altitude of these are common to the Adelbert region species. For comparison, in the mossy forest and the Huon Peninsula, and for the most at 2000-2200 m. on Mt. Kaindi, about 120 part constitute widespread forms of lowland km. south-southwest of similar elevations in and foothill habitats. Four apparently dis- the Saruwaged Range, one encounters sev- junct forms are mapped in figure 14. Several eral microhylids apparently limited to high among the remainder are yet to be resolved elevations: Barygenys flavigularis Zweifel, taxonomically. Cophixalus cryptotympanum Zweifel, C. A peculiar feature of the herpetofauna of kaindiensis Zweifel, C. parkeri Loveridge, the Adelbert Mountains is the presence there C. sp., and Phrynomantis stictogaster Zwei- of a number of species absent from the Huon fel. None of these has been found on the Pen- Peninsula, so far as is known, but recorded insula, and only C. kaindiensis and its sibling in coastal ranges 300 km. or more to the west C. parkeri have a possibly close relative of the Adelberts. The group includes two liz- there (C. biroi). It is unlikely that inadequate ards and five microhylid frogs: Cyrtodactylus collecting is the explanation. Hobart Van sermowaiensis (de Rooij) and Lobulia bron- Deusen spent the month of August 1964 at gersmai (Zweifel); Choerophryne rostellifer the "Plains of Ulur Camp" at 2380 m. in the (Wandolleck), Sphenophryne cornuta Peters Cromwell Mountains (Van Deusen, 1978) and Doria, Sphenophryne schlagenhaufeni and ascended nearby peaks to 2800 m. Ex- Wandolleck, Xenobatrachus obesus Zweifel, perienced in collecting frogs in high altitude and Xenorhina oxycephala (Schlegel). forest on Mt. Kaindi, Mt. Wilhelm, and other Collecting along the north flank of the Fin- Papuan peaks, he was astonished at the lack isterre Range adjacent to the Adelbert Moun- of frogs. In a letter written August 12, he tains may alter the picture, but at present the mentioned "have not heard a frog here." On short distance between the Adelberts and the August 24 he commented: "This gives us a Finisterres appears zoogeographically great- record of rain every day except one since we er than the much larger gap between the arrived here! . . . We have not heard a frog Adelberts and the other north coast ranges. in the area! Incredible but true." Because the Huon Peninsula region is geo- logically related to the north coast ranges of FAUNAL RELATIONSHIP TO New Guinea, it is appropriate briefly to com- NEW BRITAIN pare the faunas of the Huon Peninsula and Because of the proximity of New Britain the Adelbert Mountains, the coastal range to the Huon Peninsula, and especially be- closest to the Finisterre-Saruwaged moun- cause of the geologic relationship of the two tain mass. The Adelberts are a relatively low areas as part of the same island arc system, range, maximum elevation about 1500 m. comparison of their faunas is called for. It is Foothills of the two upland areas lie only at once evident (fig. 14) that the areas share about 30 km. apart, with the Gogol River in no highland component, although western the intervening lowlands. New Britain has elevations in excess of 1900 Information on the herpetofauna of the m. Two species of frogs common to the two Adelberts is relatively limited, but I and oth- faunas-Litoria infrafrenata and L. thesau- er members of the Alpha Helix Expedition rensis-are abundant, widespread, lowland collected there in 1969, the American Mu- forms. The single Rana known on New Brit- seum of Natural History has a small collec- ain probably falls in the same category, but tion made by E. Thomas Gilliard in 1959, and the systematics of that genus remain to be a collection of frogs, including specimens elucidated. The highly endemic frog fauna of from high elevations, is at my disposal New Britain is composed mostly of ranids thanks to Dr. Alan Ziegler of the Bishop whose relatives are in the Solomon Islands Museum. I have records of 60 species (31 to the east rather than in New Guinea. In lizards, 29 frogs) from the Adelberts and the contrast to the frogs, there are many species adjacent coastal region. At least two-thirds of lizards common to the two areas (Hediger, 1980 ZWEIFEL: HUON PENINSULA FROGS AND LIZARDS 429

1934, pp. 563-564), but these are lowland species: Cogger, 1966), almost certainly nev- forms, many of which have demonstrated er with a dry-land connection to the main- their potential for overwater dispersal by col- land. Establishment of frog populations on onizing volcanic islands such as Karkar. In the new island(s) went forward more slowly short, the frogs and lizards show no special (Karkar has only four species of frogs). In- faunal relationship between New Britain and terruption of the waterway as the Ramu- the Huon Peninsula. Faunal similarities Markham Valley rose above sea level per- among the lizards can reasonably be attrib- mitted further invasion by lizard and frog uted to overwater dispersal, and the absence species living in low, coastal habitats, merg- from New Britain of virtually all of the lowing the insular, waif fauna with the more di- land frog component of the Peninsula speaks verse and widespread lowland assemblage of for the lack of any previous land connection the New Guinea mainland. Elements of this between the two regions. coastal fauna invaded the uplands in varying degree, but higher elevations had relatively HISTORY OF THE PENINSULAR meager faunas. The coming of Pleistocene FAUNA glaciation, with concurrent cooling and lowered sea level, changed the climate of the The following points may be gleaned from Ramu-Markham Valley sufficiently to permit the preceding discussion: (1) the block of a variety of lower montane species of frogs land of which the Huon Peninsula is part and lizards to disperse across the Valley and underwent most of its uplift since the late become established before changing climate Pliocene and from its origin through part of again made the Valley unsuitable. Never did its history was an island, separated from the the climate change sufficiently to permit mainland of New Guinea by an arm of the forms now restricted to high, cool elevations sea along the axis of the present Ramu-Mark- of the major New Guinea mountains to make ham Valley; (2) endemism among frogs is so the crossing. low as to be virtually negligible and is non- Thus, the present-day frog and existent at the species level among lizards; lizard fau- (3) 48 of 52 species of lizards in the study nas of the Huon Peninsula are seen to be area (92%) and 19 of 35 nonendemic species composed of two principal elements: low- of native frogs (54%) inhabit lowlands: (4) land species, many of them widespread, with among the species of frogs and lizards with access (in a genetic sense) to populations disjunct populations on the Peninsula, all outside the Peninsula; and a minority of range at least as low as 1400 m., most to 1100 species occupying higher elevations and m., and some even below 1000 m.; (5) the presently cut off from conspecific popula- Peninsula appears to lack a high altitude frog tions off the Peninsular uplands. For the dis- fauna such as is present at similar elevations junct species, the stage is set for allopatric along the central montane axis of New speciation events, but the degree of ende- Guinea. mism detected so far is minimal. In this in- Given the foregoing observations, the fol- stance, geologic evolution seems to have lowing hypothetical history of the frog and outpaced organic. lizard faunas is advanced. The first lands of These hypotheses of the derivation of the the Finisterre-Saruwaged mass rising above fauna may be testable by comparison with the tropical sea in the Pliocene were popu- other animal groups with similar dispersal lated by a variety of lizards, mostly skinks potentials. If these prove to have high de- but some geckos, dispersing across the rel- grees of endemism and well-developed high atively narrow waterway from larger land montane faunas, then one must look to other masses to the south and west. The potential views of distributional and geological history for such dispersal seems high, in view of the or to vastly different rates of speciation for relatively diverse reptile faunas of small vol- alternative explanations. canic islands such as Karkar (about 30 430 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 165

LITERATURE CITED Barker, John, and Gordon Grigg Brown, Walter C., and Fred Parker 1977. A field guide to Australian frogs. Ade- 1977. Lizards of the genus Lepidodactylus laide, Rigby Ltd., pp. 1-229, figs., pls., (Gekkonidae) from the Indo-Australian maps. Archipelago and the islands of the Pa- Boulenger, G. A. cific, with descriptions of new species. 1887. Catalogue of the lizards in the British Proc. California Acad. Sci., 4th ser., Museum (Natural History). London, vol. 41, no. 8, pp. 253-265, 1 fig., 1 ta- vol. 3, xii + 575 pp., 40 pls. ble. 1897. Descriptions of new lizards and frogs Bulmer, R. N. H., and M. J. Tyler from Mount Victoria, Owen Stanley 1968. Karam classification of frogs. Jour. Range, New Guinea, collected by Mr. Polynesian Soc., vol. 77, pp. 333-385, A. S. Anthony. Ann. Mag. Nat. Hist., table 1. ser. 6, vol. 19, pp. 6-13, pls. 1-2. Burt, Charles E., and May D. Burt 1898. An account of the reptiles and batrachi- 1932. Herpetological results of the Whitney ans collected by Dr. L. Loria in British South Sea Expedition. VI. Pacific island New Guinea. Ann. Mus. Civ. Stor. Nat. amphibians and reptiles in the collection Genova, ser. 2, vol. 18, pp. 694-710, of the American Museum of Natural pls. 6-8. History. Bull. Amer. Mus. Nat. Hist., 1903. Descriptions of new reptiles from Brit- vol. 63, art. 5, pp. 461-597, figs. 1-38. ish New Guinea. Proc. Zool. Soc. Lon- Chappell, John don, vol. 2, pp. 125-129, pls. 12-13. 1974. Geology of coral terraces, Huon Pen- insula, New Guinea: a study of Quater- Brongersma, L. D. nary tectonic movements and sea-level 1933. A new gecko of the genus Gymnodac- changes. Geol. Soc. Amer. Bull., vol. tylus from New Guinea. Ann. Mag. Nat. 85, pp. 553-570, figs. 1-20, table 1. Hist., ser. 10, vol. 11, pp. 252-253. Cogger, Harold C. 1934. Contributions to Indo-Australian her- 1975. Reptiles and amphibians of Australia. petology. Zool. Meded., vol. 17, pp. i- Sydney, A. H. and A. W. Reed, 585 pp., xvi, 161-251, figs. 1-47, pls. 1-2. figs., maps. 1949. Zoological results of the Dutch New Costin, Alec B., Ru D. Hoogland, and C. Lendon Guinea Expedition, 1939. No. I. A new 1977. Vegetation changes in the Lake Mamsin scincid lizard. Nova Guinea, n. ser., area, Saruwaged Plateau, New Guinea. vol. 5, pp. 272-283, figs. 1-4, pls. 1-2. Amer. Mus. Novitates, no. 2628, pp. 1- 1953. Notes on New Guinean reptiles and am- 9, figs. 1-4, tables 1-3, map 1. phibians III. Proc. K. Nederl. Akad. Dessauer, Herbert C., Donald F. Gartside, and Wetensch., ser. C, vol. 56, pp. 572-583, Richard G. Zweifel figs. 1-2. 1977. Protein electrophoresis and the system- Brown, Walter C. atics of some New Guinea hylid frogs 1953. Results of the Archbold Expeditions. (genus Litoria). Syst. Zool., vol. 26, no. No. 69. A review of New Guinea lizards 4, pp. 426-436, figs. 1-2, tables 1-3. allied to Emoia baudini and Emoia Dow, D. B. physicae (Scincidae). Amer. Mus. Nov- 1977. A geological synthesis of Papua New itates, no. 1627, pp. 1-25, figs. 1-6, ta- Guinea. Dept. Natl. Resources Bur. bles 1-4. Min. Resources Geol. Geophys. Bull., 1954. Notes on several lizards of the genus no. 201, pp. i-vi, 1-41, figs. 1-36, pls. Emoia with descriptions of new species (maps) 1-8. Canberra, Australian Govt. from the Solomon Islands. Fieldiana: Publ. Serv. Zool., vol. 34, no. 25, pp. 263-276, figs. Duellman, William E. 44-45, tables 1-2. 1967. Additional studies of chromosomes of 1956. The distribution of terrestrial reptiles in anuran amphibians. Syst. Zool., vol. 16, the islands of the Pacific basin. Proc. no. 1, pp. 38-43, figs. 1-8. 8th Pacific Sci. Congr., vol. 3 A, Ocean- Forcart, Lothar ogr. Zool., pp. 1479-1491, tables 1-4. 1953. Amphibien und Reptilien von Neugui- 1980 ZWEIFEL: HUON PENINSULA FROGS AND LIZARDS 431

nea, mit der Beschreibung eines neuen morphus variegatus (Peters). Fieldiana: Laubfrosches, Nyctimystes flavomacu- Zool., vol. 39, no. 24, pp. 257-268, figs. lata n. sp. Verh. Naturf. Ges. Basel, 41-45, table 1. vol. 64, no. 1, pp. 58-68, fig. 1. 1961. Notes on two New Guinean lizards of Greer, Allen E. the genus Sphenomorphus. Ibid., vol. 1968. Clutch size in the scincid genus Emoia. 39, no. 47, pp. 539-542, fig. 102, table 1. Copeia, no. 2, pp. 417-418, fig. 1, ta- Jaques, A. L., and G. P. Robinson ble 1. 1977. The continent/island arc collision in 1970a. A subfamilial classification of scincid northern Papua New Guinea. BMR lizards. Bull. Mus. Comp. Zool., vol. Jour. Australian Geol. Geophys., vol. 139, no. 3, pp. 151-183, figs. 1-7. 21, no. 4, pp. 289-303, figs. 1-9, ta- 1970b. The relationships of the skinks referred ble 1. to the genus Dasia. Breviora, no. 348, Jong, J. K. de pp. 1-30, figs. 1-6, tables 1-2. 1927. Reptiles from Dutch New Guinea. Nova 1973. Two new lygosomine skinks from New Guinea, vol. 15, no. 3, pp. 296-318, figs. Guinea with comments on the loss of 1-4. the external ear in lygosomines and ob- Kampen, P. N. van servations on previously described 1913. Amphibien, gesammelt von der Nied- species. Ibid., no. 406, pp. 1-25, figs. 1- erliindischen Sud.-Neu-Guinea-Expedi- 10, tables 1-2. tion von 1909-10. Nova Guinea, vol. 9, 1974. The generic relationships of the scincid Zool., pt. 1, pp. 453-465, pl. 11. lizard genus Leiolopisma and its rela- Kluge, Arnold G. tives. Australian Jour. Zool., suppl. 1974. A taxonomic revision of the lizard fam- ser., no. 31, pp. 1-67, figs. 1-64. ily Pygopodidae. Misc. Publ. Mus. Greer, Allen E., and Fred Parker Zool. Univ. Michigan, no. 147, pp. i-vi, 1974. The fasciatus species group of Sphen- 1-221, figs. 1-124, tables 1-22. omorphus (Lacertilia: Scincidae): notes Kopstein, Felix on eight previously described species 1926. Reptilien von den Molukken und den and descriptions of three new species. benachbarten Inseln. Zool. Meded., Papua New Guinea Sci. Soc. Proc., vol. vol. 9, pp. 71-112. 25, pp. 31-61. Loeffler, R. Greer, Allen E., and Gary Raizes 1971. The Pleistocene glaciation of the Saru- 1969. Green blood pigment in lizards. Sci- waged Range, Territory of New Guinea. ence, vol. 166, no. 3903, pp. 392-393. Australian Geogr., vol. 11, no. 5, pp. Hediger, Heini 463-472, figs. 1-2, pls. 1-6. 1934. Beitrag zur Herpetologie und Zoogeo- 1972. Pleistocene glaciation in Papua New graphie New Britanniens und einiger Guinea. Zeitschr. Geomorph., suppl. umliegender Gebiete. Zool. Jahrb. vol. 13, pp. 32-58, figs. 1-8, table 1. (Syst.), vol. 65, nos. 5-6, pp. 389-582, Lonnberg, Einar figs. 1-6. 1900. Reptiles and batrachians collected in Hope, G. S., and J. A. Peterson German New Guinea by the late Dr. 1975. Glaciation and vegetation in the high Erik Nyman. Ann. Mag. Nat. Hist., ser. New Guinea mountains. In Suggate, R. 7, vol. 6, pp. 574-582. P., and M. M. Cresswell (eds.), Quater- Loveridge, Arthur nary studies. Wellington, Royal Soc. 1945. New scincid lizards of the genera Tro- New Zealand, pp. 155-162, figs. 1-6. pidophorus and Lygosoma from New Hope, J. H., and G. S. Hope Guinea. Proc. Biol. Soc. Washington, 1976. Palaeoenvironments for man in New vol. 58, pp. 47-5 1. Guinea. In Kirk, R. L., and A. G. 1948. New Guinean reptiles and amphibians Thorne (eds.), The origin of the Austra- in the Museum of Comparative Zoology lians. Canberra, Australian Inst. Ab- and United States National Museum. original Studies, pp. 29-54, figs. 1-4, Bull. Mus. Comp. Zool., vol. 101, no. tables 1-2. 2, pp. 305-430. Inger, Robert F. Maderson, P. F. A. 1958. Three new skinks related to Spheno- 1970. Lizard glands and lizard hands: models 432 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 165

for evolutionary study. Forma Functio, 1936. A collection of reptiles and amphibians vol. 3, pp. 179-204, figs. 1-16, tables from the mountains of British New 1-2. Guinea. Ann. Mag. Nat. Hist., ser. 10, Mehely, L. von vol. 17, pp. 66-93, figs. 1-6. 1897. Ujabb adatok Uj-Guinea herpetologia- Peters, W., and G. Doria jahoz. [Further contributions to the her- 1878. Catalogo dei Rettili e dei Batraci rac- petology of New-Guinea.] Termesz. colti da 0. Beccari, L. M. D'Albertis e Fiiz., vol. 20, pp. 398-419, 1 pl. (Hun- A. A. Bruijn nella sotto-regione Austro- garian text, pp. 398-409, followed by Malese. Ann. Mus. Civ. Stor. Nat. Gen- English translation.) ova, vol. 13, pp. 325-450, figs., pls. 1- 1901. Beitrage zur Kenntniss der Engysto- 7, tables. matiden von Neu-Guinea. Ibid., vol. 24, Rooij, Nelly de pp. 169-271, pls. 4-12. (Hungarian text, 1915. The reptiles of the Indo-Australian Ar- pp. 169-216; German translation, pp. chipelago. I. Lacertilia, Chelonia, Emy- 216-267.) dosauria. Leiden, E. J. Brill, Ltd., pp. Menzies, J. I. i-xiv, 1-384, figs. 1-132. 1972. Papuan tree frogs of the Litoria nigro- 1919. Reptilien aus Nord-Neuguinea gesam- punctata group. Herpetologica, vol. 28, melt von Dr. P. N. van Kampen und Dr. no. 4, pp. 291-300, figs. 1-5, tables 1-2. K. Gjellerup in den Jahren 1910 und 1976. Handbook of common New Guinea 1911. Bijd. Dierk., vol. 21, pp. 81-95, frogs. Wau Ecology Institute, Hand- figs. 1-10. book no. 1, viii + 75 pp., figs. 1-2, pls. Smith, Malcolm A. 1-12. 1937. A review of the genus Lygosoma (Scin- Menzies, J. I., and Michael J. Tyler cidae: Reptilia) and its allies. Rec. In- 1977. The systematics and adaptations of dian Mus., vol. 39, pt. 3, pp. 213-234, some Papuan microhylid frogs which figs. 1-5. live underground. Jour. Zool. (Lon- Tyler, Michael J. don), vol. 183, pp. 431-464, figs. 1-14, 1963a. A taxonomic study of amphibians and pls. 1-3, tables 1-2. reptiles of the Central Highlands of New Menzies, J. I., and George R. Zug Guinea, with notes on their ecology and 1979. Papuan tree frogs of the Litoria thesau- biology. I. Anura: Microhylidae. Trans. rensis group (Salientia: Hylidae). Mi- Roy. Soc. South Australia, vol. 86, pp. cronesica, vol. 15, nos. 1-2, pp. 325- 11-29, figs. 1-2, table 1. 333. 1963b. A taxonomic study of amphibians and Menzies, J. I., and Richard G. Zweifel reptiles of the Central Highlands of New 1974. Systematics ofLitoria arfakiana of New Guinea, with notes on their ecology and Guinea and sibling species (Salientia, biology. II. Anura: Ranidae and Hyli- Hylidae). Amer. Mus. Novitates, no. dae. Ibid., vol. 86, pp. 105-130, figs. 1- 2558, pp. 1-16, figs. 1-8, tables 1-2. 7, tables 1-2. Mertens, Robert 1963c. An account of collections of frogs from 1931. Ablepharus boutonii (Desjardin) und Central New Guinea. Rec. Australian seine geographische Variation. Zool. Mus., vol. 26, no. 3, pp. 113-130, figs. Jahrb., vol. 61, nos. 1-2, pp. 63-210, 1-2, pls. 1-5, table 1. figs. 1-6, pls. 2-4. 1964a. Transfer of the New Britain frog, Hyla 1942. Die Familie der Warane (Varanidae). brachypus (Werner) to the microhylid III: Taxonomie. Abhandl. Senckenberg. genus Oreophryne. Mitt. Zool. Mus. Naturf. Gesell., no. 466, pp. 235-391. Berlin, vol. 40, no. 1, pp. 3-8, figs. 1-3, Mittleman, M. B. table 1. 1952. A generic synopsis of the lizards of the 1964b. An investigation of the systematic po- subfamily Lygosominae. Smithsonian sition and synonymy of Hyla montana Misc. Coll., vol. 117, no. 17, pp. 1-35. Peters and Doria (Anura, Hylidae). Parker, H. W. Zool. Abhandl. Mus. Tierk. Dresden, 1934. A monograph of the frogs of the family vol. 27, no. 10, pp. 265-270, 2 figs., 1 Microhylidae. London, British Museum table. (Natural History), viii + 208 pp., figs. 1965. Taxonomic studies of some hylid frogs 1-67, 2 maps. of Australia and New Guinea. Proc. 1980 ZWEIFEL: HUON PENINSULA FROGS AND LIZARDS 433

Zool. Soc. London, vol. 145, pt. 1, pp. Bismarck-Schrader Range, New Guinea. 91-106, figs. 1-5, pl. 1, tables 1-3. Mem. Nat. Mus. Victoria, vol. 33, pp. 1967. Microhylid frogs of New Britain. Trans. 57-63, fig. 1, pl. 8. Roy. Soc. South Australia, vol. 91, pp. 187-190. Zug, George R., Eric Lindgren, and John R. Pip- 1968. Papuan hylid frogs of the genus Hyla. pet Zool. Verh. Leiden, no. 96, pp. 1-203, 1975. Distribution and ecology of the marine figs. 1-58, pls. 1-4, tables 1-2. toad, Bufo marinus, in Papua New 1971. The phylogenetic significance of vocal Guinea. Pacific Sci., vol. 29, no. 1, pp. sac structure in hylid frogs. Univ. Kan- 31-50, figs. 1-5, tables 1-6. sas Publ. Mus. Nat. Hist., vol. 19, no. Zweifel, Richard G. 4, pp. 319-360, figs. 1-10, tables 1-3. 1956a. Results of the Archbold Expeditions. 1972. An analysis of the lower vertebrate fau- No. 72. Microhylid frogs from New nal relationships of Australia and New Guinea, with descriptions of new Guinea. In Walker, D. (ed.), Bridge and species. Amer. Mus. Novitates, no. barrier: the natural and cultural history 1766, pp. 1-49, figs. 1-9, tables 1-4. of Torres Strait. Australian Natl. Univ. 1956b. Notes on microhylid frogs, genus Co- Res. School Pacific Studies Dept. Bio- phixalus, from New Guinea. Ibid., no. geogr. Geomorph. Publ. BG/3, pp. 231- 1785, pp. 1-8, figs. 1-3. 256. 1958. Results of the Archbold Expeditions. Tyler, Michael J., and Margaret Davies No. 78. Frogs of the Papuan hylid genus 1978. Species-groups within the Australopa- Nyctimystes. Ibid., no. 1896, pp. 1-51, puan hylid frog genus Litoria. Austra- figs. 1-21, tables 1-2. lian Jour. Zool., suppl. ser. no. 63, pp. 1962. Results of the Archbold Expeditions. 1-47, figs. 1-27. No. 83. Frogs of the microhylid genus Tyler, Michael J., and J. I. Menzies Cophixalus from the mountains of New 1971. A new species of microhylid frog of the Guinea. Ibid., no. 2087, pp. 1-26, figs. genus Sphenophryne from Milne Bay, 1-7, tables 1-2. Papua. Trans. Roy. Soc. South Austra- 1966. A new lizard of the genus Tribolonotus lia, vol. 95, pt. 2, pp. 79-83, figs. 1-2. (Scincidae) from New Britain. Ibid., no. Van Deusen, Hobart M. 2264, pp. 1-12, figs. 1-3, table 1. 1978. Results of the Archbold Expeditions. 1969. Frogs of the genus Platymantis (Ran- No. 101. Summary of the Seventh Arch- idae) in New Guinea, with the descrip- bold Expedition to New Guinea (1964). tion of a new species. Ibid., no. 2374, Amer. Mus. Novitates, no. 2660, pp. 1- pp. 1-19, figs. 1-5, table 1. 21, figs. 1-12. 1972a. Results of the Archbold Expeditions. Vogt, Theodor No. 97. A revision of the frogs of the 1911. Reptilien und Amphibien aus Neu- subfamily Asterophryinae, family Mi- Guinea. Sitzber. Gesell. Naturf. Freunde crohylidae. Bull. Amer. Mus. Nat. Berlin, no. 9, pp. 410-420. Hist., vol. 148, art. 3, pp. 411-546, figs. 1912. Reptilien und Amphibien aus Hollan- 1-79, tables 1-9. disch-Neu-Guinea. Ibid., no. 6, pp. 355- 1972b. A new scincid lizard of the genus Leio- 359. lopisma from New Guinea. Zool. Med- 1932. Beitrag zur Reptilienfauna der ehema- ed., vol. 47, pp. 530-539, figs. 1-3, pl. 1. ligen Kolonie Deutsch-Neuguinea. Ibid., 1972c. A review of the frog genus Lechriodus nos. 5-7, pp. 281-294. (Leptodactylidae) of New Guinea and Wermuth, Heinz Australia. Amer. Mus. Novitates, no. 1965. Liste der rezentan Amphibien und Rep- 2507, pp. 1-41, figs. 1-13, table 1. tilien. Gekkonidae, Pygopodidae, Xan- 1972d. Frogs. In Ryan, Peter (ed.), Encyclo- tusiidae. Das Tierreich, no. 80, pp. i- paedia of Papua New Guinea, vol. 1, A- xxii, 1-246. K. Melbourne, Univ. Press, pp. 466- 1967. Liste der rezenten Amphibien und Rep- 471. tilien. Agamidae. Ibid., no. 86, pp. xiv + 1976. Herpetological expedition to New 127. Guinea. Natl. Geogr. Soc. Res. Repts., Woodruff, David S. 1968 Projects, pp. 503-510, 1 fig. 1972. Amphibians and reptiles from Simbai, 1977. Rokrok. Things that go croak in the 434 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 165

night. Animal Kingdom, April/May, pp. frog (genus Cophixalus) from Papua 28-34, figs. New Guinea, with notes on related 1979a. Variation in the scincid lizard Lipinia forms. Ibid., no. 2678, pp. 1-14, figs. noctua and notes on other Lipinia from 1-8. the New Guinea region. Amer. Mus. Zweifel, Richard G., and Fred Parker Novitates, no. 2676, pp. 1-21, figs. 1- 1977. A new species of frog from Australia 12. (Microhylidae, Cophixalus). Amer. Mus. 1979b. A new cryptic species of microhylid Novitates, no. 2614, pp. 1-10, figs. 1-7. BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY

Volume 165 1979-1980

PUBLISHED BY ORDER OF THE TRUSTEES NEW YORK: 1980 Edited by FLORENCE BRAUNER CONTENTS OF VOLUME 165

Article 1. The Systematics, Phylogeny, and Zoogeography of the Blissinae of the World (Hemiptera, Lygaeidae). By James A. Slater. Pages 1-180, figures 1-80. De- cember 27, 1979 ...... Price. $11.60 Article 2. Calmoniid Trilobites of the Lower Devonian Scaphiocoelia Zone of Bolivia, With Remarks on Related Species. By Niles Eldredge and Leonardo Branisa. Pages 181-290, figures 1-38, tables 1, 2. June 13, 1980 ...... Price. $6.95 Article 3. A Revision of the Moth Genus Somatolophia (Lepidoptera, Geometridae). By Frederick H. Rindge. Pages 291-334, figures 1-59, maps 1-3, table 1. July 11, 1980 ...... Price. $2.75 Article 4. A Revision of the Spider Genus Cesonia (Araneae, Gnaphosidae). By Norman I. Platnick and Mohammed U. Shadab. Pages 335-386, figures 1-145, maps 1-6. July 11, 1980 ...... Price. $3.25 Article 5. Results of the Archbold Expeditions. No. 103. Frogs and Lizards from the Huon Peninsula, Papua New Guinea. By Richard G. Zweifel. Pages 387-434, figures 1-14, 1 table. September 22, 1980 ...... Price. $3.10